(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Biodiversity Heritage Library | Children's Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Proceedings of the Linnean Society of New South Wales"

PROCEEDINGS 

of the 



LINNEAN 
SOCIETY 



NEW SOUTH WALES 




NATURAL HISTORY IN ALL ITS BRANCHES 



THE LINNEAN SOCIETY OF 

NEW SOUTH WALES 

ISSN 0370-047X 




Founded 1874 
Incorporated 1 884 

The Society exists to promote the cultivation and study 
of the science of natural history in all its branches. 
The Society awards research grants each year in the 
fields of Life Sciences (the Joyce Vickery fund) and 
Earth Sciences (the Betty Mayne fund), offers annually 
a Linnean Macleay Fellowship for research, contributes 
to the stipend of the Linnean Macleay Lecturer in 
Microbiology at the University of Sydney, and 
publishes the Proceedings. It holds field excursion and 
scientific meetings, including the biennial Sir William 
Macleay Memorial Lecture delivered by a person 
eminent in some branch of natural science. 



Membership enquiries should be addressed in the first instance to the Secretary. Candidates for elec- 
tion to the Society must be recommended by two members. The present annual subscription is 
$A53.00. 

The current subscription rate to the Proceedings is set at $A80.00 per volume. In recent years a 
volume consists of a single annual issue. 

Back issues of all but a few volumes and parts of the Proceedings are available for purchase. Prices 
are listed on our home page and can also be obtained from the Secretary. 

OFFICERS AND COUNCIL 2003/2004 



President: I.G. Percival 

Vice-presidents: R.J. King, K.L. Wilson, A. Ritchie, J.R Barkas 

Honorary Treasurer: M.L. Augee 

Secretary: J-C. Herremahs 

Council: A.E.J. Andrews, M.L. Augee, J.R Barkas, M.R. Gray, J-C. Herremans, M.A. Humphrey, 

D. Keith, R.J. King, H.A. Martin, P.M. Martin, J.R. Merrick, M.S. Moulds, D.R. Murray, P.J. 

Myerscough, I.G. Percival, A. Ritchie, S. Rose, and K.L. Wilson 

Honorary Editor: M.L. Augee 

Linnean Macleay Lecturer in Microbiology: PR. Reeves 

Auditors: Phil Williams Carbonara 

The postal address of the Society is: P.O. Box 82, Kingsford NSW 2032, Australia 

Telephone: (International) 61 2 9662 6196; (Aust) 02 9662 6196 

E-mail: linnsoc@acay.com.au 

Home page: www.acay.com.au/~linnsoc/welcome.html 

© Linnean Society of New South Wales 

Cover motif: diving platypus by Marianne Larsen, Wellington NSW. 






PROCEEDINGS 11 MM 2004 

LINNEAN 
SOCIETY 

of 

NEW SOUTH WALES 



For information about the Linnean Society of New South Wales, 
its publications and activities, see the Society's homepage 

http://www.acay.com.au/~linnsoc/welcome.htm 



VOLUME 125 

February 2004 



EDITORIAL 



This volume consists of three parts. The first part contains general contributions. The second part contains 
research papers and review papers arising from the symposium "Monotreme HI" held by the Linnean Society of 
NSW and the Australian Mammal Society at the University of Sydney in July 2003. The third section contains 
book reviews and an obituary to Merv Griffiths, the pre-eminent monotreme biologist of all time. 

The publication of this volume has been delayed by the preparation of the papers from "Monotreme III" and is 
covered by subscriptions and membership fees for 2003. 

Intending authors should read the summary of "Instructions for Authors" at the back of this volume carefully. 
More details are available in the full version available at the Society's web site or from the Secretary. The 
preparation of this volume has been prolonged and made difficult by the failure of some authors to provide 
figures and tables in the format required. In order to keep the costs at a level which allows our Society to 
continue publication, we set the journal completely ourselves. Therefore we do not have the flexibility of large 
commercial publishers. We can only deal with figures as photographs, original line drawings or .TEF files. Jpeg 
files for example are useless in our system. Auto-formatting and track changes are a disaster, as are tables and/ 
or figures that have been put inside the text. To date we have taken the time to re-set and sometimes re-scan 
figures, however in future we will apply the policy that final copy not prepared in accordance with the instructions 
will simply be returned and held over if necessary until the next issue. 



M.L. Augee 
Editor 



Review of Australian Cave Guano Ecosystems with a Checklist 

of Guano Invertebrates 

Timothy Moulds 

Centre for Evolutionary Biology and Biodiversity, School of Earth and Environmental Sciences, University 

of Adelaide, North Terrace, Adelaide 5005. 
timothy.moulds @ adelaide.edu.au 



Moulds, T. (2004). Review of Australian cave guano ecosystems with a checklist of guano invertebrates. 
Proceedings of the Linnean Society of New South Wales 125, 1-42. 

This work provides a check-list of all invertebrate species known, or believed to be, associated 
with cave guano in Australia. A total of 240 species in 121 families, representing 25 orders is listed. These 
species inhabit 60 karst areas in all mainland states of Australia and Christmas Island (Indian Ocean). 
Comprehensive assessment of all available records (published and in collections) show that the distribution 
of several species is more extensive than previously believed. It is unknown whether this is because of 
inadequate identification of specimens, poorly defined taxonomy or unrecognised intra-species variation 
due to a lack of specimens. Twenty species from five orders show restricted distributions and guano 
dependence, although endemic status can not yet be assigned. Amongst these species, eight pseudoscorpions 
and eight Coleoptera are distributed across several mainland states and Christmas Island. 

Manuscript received 2 July 2003, accepted for publication 22 October 2003. 

Keywords: Arthropoda, Australia, biospeleology, cave, checklist, ecosystem, guano, invertebrate. 



INTRODUCTION 

Australian cave guano ecosystems are poorly 
known with only a few communities studied in detail 
(e.g. Richards 1971; Harris 1973; Bellati et al. 2003). 
Previous studies concerned with the terrestrial 
cavernicolous fauna of Australia have mentioned 
species associated with guano, but have provided little 
in the way of detail with regard to the ecology of 
specialised guano communities and species. This paper 
seeks to synthesise the knowledge of guano ecosystems 
and communities in Australian caves. A review of 
guano ecosystems and habitats precedes a checklist of 
all species known to be associated with Australian cave 
guano deposits. 

Populations of cavernicolous animals are 
usually small because of limited food supplies. 
However, caves containing guano differ fundamentally 
because there is a virtually unlimited food supply, 
commonly resulting in large animal populations. 
Guano in caves is deposited by bats, birds, orthopterans 
(crickets and grasshoppers), and small mammals, with 
each type of guano sustaining a unique assemblage of 
taxa. Guano deposits are extremely variable, unlike 
other cave habitats, and consist of numerous micro- 



habitats differentiated by fluctuating temperature, 
moisture, and pH. Guano ecosystems contain obligate 
guano-dwelling organisms (guanobites), opportunistic 
guano-dwelling animals (guanophiles), and transient 
guano-using animals (guanoxenes) (Gnaspini and 
Trajano 2000). The basis for many guano ecosystems 
is the numerous species of fungi and bacteria that can 
grow on guano, even in complete darkness. 

Cave food sources 

Cavernicolous populations are dependant for 
their survival upon energy inputs into cave systems. 
These inputs can vary widely, with availability of food 
usually being the primary limiting factor (Peck 1976). 
Inflowing streams and periodic floodwaters introduce 
significant amounts of zooplankton, accidentals, and 
organic debris that, for many cave ecosystems, 
represent the main energy inputs (Peck and 
Christiansen 1990; Humphreys 1991). Tree roots 
penetrating the roofs and walls are another important 
energy source found commonly in tropical caves and 
lava tubes (Hoch 1988; Hoch and Howarth 1999). 
Dead animals can be a source of food for scavengers 
near cave entrances (Richards 1971). Accidentals 
wandering in from cave entrances also provide a food 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



source, although this is generally periodic in nature 
and inconsistent in quantity. 

For the most part cave environments are 
generally depaurporate in food and consequently are 
sparsely populated with cavernicolous animals. 
However, caves containing guano deposits differ 
substantially because they have a virtually unlimited 
food supply. When present, guano from bats, birds, 
and Orthoptera (crickets and grasshoppers) generally 
forms the major energy source (Park and Barr 1961; 
Poulson 1972; Martin 1977), with large, varied and 
unique ecosystems often revolving around such 
deposits. 



SOURCES AND DIVERSITY OF CAVE GUANO 

Cave guano deposits from specific sources 
can each possess a unique assemblage of taxa (Horst 
1972; Poulson 1972). Throughout the world's 
biogeographic provinces different taxa are responsible 
for being the most important guano producers. The 
most widespread and common guano is that produced 
by bats and these deposits are generally the largest in 
volume. The spatial and temporal deposition of bat 
guano differs from tropical to temperate caves. Cave- 
dwelling bats in temperate regions show an annual 
cycle of occupancy over summer months when pups 
are born, before colonies disperse to cooler, wintering 
caves where they enter torpor. This annual cycle results 
in large amounts of guano deposited over summer 
months and then a cessation of guano input for 
approximately seven months. In contrast, tropical caves 
generally show constant bat occupancy rather than an 
annual cycle and less congregation of individuals due 
to warmer ambient temperatures. Gnaspini and Trajano 
(2000) note that many bat populations in tropical Brazil 
are commonly nomadic, resulting in roaming colonies 
varying their location in an irregular and non-seasonal 
fashion. This results in non-continuous deposition. The 
diet of bats (either haematophagous, insectivorous, 
frugivorous, or nectarivorous) also influences the 
composition of guano piles and hence the associated 
guanophilic communities (Gnaspini 1992; Ferreira and 
Martins 1998, 1999). Large populations of the common 
vampire bat (Desmodus rotundus Geoffroy) 
predominate in Brazilian karst near inhabited areas, 
due to large numbers of domestic livestock resulting 
in haematophagous guano deposits. Guano from non- 
haematophagous bats is absent, or greatly reduced as 
vampire bats exclude other bat species, thus changing 
the guanophilic communities present. 

Birds are common guano producers in the 
northern parts of South America, the Caribbean and 



tropical caves of south-east Asia. Cave-dwelling birds 
nest in the dark zone, providing an important energy 
resource for many cavernicolous animals. Cave- 
dwelling birds in South American and Caribbean caves 
include guacharos (Steatornis caripensis Humboldt) 
(Snow 1975; Gnaspini and Trajano 2000). This bird 
discards palm seeds, sometimes with flesh still 
attached, and deposit droppings in caves, thus 
providing a wide range of organic matter for 
cavernicolous animals. Because of the presence of 
discarded seeds, some taxa associated with seeds and 
detritus, such as lygaeid bugs are found only in guano 
of this type. Swiftlets (Aerodramus spp) nest in the 
entrance and dark zones of tropical caves in south- 
east Asia, northern Australia and the Pacific and are 
insectivorous (Medway 1962). These birds also support 
a range of guanophilic taxa in the caves of Christmas 
Island (Humphreys and Eberhard 2001). Richards 
(1971) reported that droppings from several species 
of birds nesting in the entrance zone of Nullarbor Plain 
caves support a wide variety of cavernicolous animals. 

Rhaphidophorid crickets are often important 
producers of guano in temperate caves such as those 
of the Nullarbor Plain (Richards 1971). The sometimes 
large populations of these crickets can accumulate 
sizeable guano deposits in caves. These deposits are 
important as few other food sources exist in areas such 
as the Nullarbor Plain because the low mean rainfall 
limits organic flood debris and bat populations are 
generally small. Rhaphidophorid guano is also utilised 
in Mammoth Cave, Kentucky, where it is widely 
dispersed through the cave system (Howarth 1983). 

Small mammals are often significant guano 
producers in temperate zones of North America. The 
guano of porcupines (Erethizon dorsatum L.) is 
reported by Calder (1965) to support a community of 
collembolans and mites active throughout the year in 
Frenchman's Cave (Hants County, Nova Scotia, 
Canada). Cave rats (Neotoma spp), navigate using 
urine trails (Howarth 1983). Although common in the 
caves of the Canadian Rockies and Vancouver Island, 
their faeces are mostly unusable as a food source due 
to the high ammonia content from systematic urination 
at these sites (Trapani 1997). 



GUANO ECOSYSTEMS AND FOOD WEBS 

Guanobites are animals that require the 
presence of guano for survival. They will only feed 
on guano and will not use other food sources within 
caves. Although guanobitic species are occasionally 
found on other substrates in caves as they move 
between discontinuous guano deposits, they do not feed 



Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



or reproduce on these substrates (Gnaspini and Trajano 
2000). When guano deposition is seasonal (e.g. bat 
maternity caves), guanobites will commonly become 
quiescent until bats return and restore fresh guano 
input. Other guanobite populations crash when guano 
input ceases and then quickly reproduce when guano 
input recommences. 

Guanophiles use guano resources 
opportunistically and are able to complete their entire 
life cycle using the guano substrate. Guanophiles will 
however utilise other cave food resources when 
available and do not have to rely upon guano to feed 
or reproduce. Abundance of guanophilic animals will 
decrease if fresh guano is not available, simply due to 
food limitation, but individuals will attempt to exploit 
other food resources to survive until fresh guano is 
available. Troglobites and troglophiles that have a 
generalist role in epigean ecosystems are classified as 
guanophiles if they utilise guano when available, even 
though they are capable of surviving subterranean 
habitats without this resource. 

Guanoxenes will exploit a guano resource for 
feeding or reproduction but require other substrates 
within a cave to complete their life cycle (Gnaspini 
and Trajano 2000). Guanoxenes can be either 
troglobites, troglophiles or trogloxenes (Gnaspini and 
Trajano 2000). 

The cyclical nature of many guano deposits 
resulting from the annual breeding cycle of bats, leads 
to a similar cycle in arthropod abundances. Low 
population numbers of many species reflect changes 
in micro-habitat conditions resulting from the cessation 
of fresh guano deposition and lower air and guano 
temperatures. Guano communities decrease in numbers 
as many species stop breeding until the food supply 
(i.e. fresh guano) is restored. This has been observed 
in the mite Uroobovella coprophila Womersley, which 
is quiescent during winter months in Carrai Bat Cave, 
northern New South Wales (Harris 1971). 

Arthropods in guano communities feed either 
directly on guano or fungus growing on guano deposits 
and these in turn support a number of predators 
scavengers and omnivores (Gillieson 1997). 
Generalised guano food webs have a guano source 
directly supporting a range of guanivores including 
Phoridae (Diptera), Anobiidae (Coleoptera), Tineidae 
(Lepidoptera), Collembola and mesostigmatid mites 
(Acarina). Predators that prey upon these consumers 
include spiders, pseudoscorpions, beetles and 
opiliones. Specialised parasites and parasitoids are also 
active in many guano ecosystems. Braconid wasps 
(Hymenoptera) are found in many Australian guano 
caves and parasitise the larvae of Monopis spp 
(Lepidoptera: Tineidae). The larvae of the guanobite 



Derolathrus sp. (Coleoptera: Jacobsoniidae) are - 
parasitised by small myrmarid wasps (Hymenoptera). 
Parasitic relationships in guano ecosystems are 
generally poorly understood and further research will 
undoubtedly reveal many more examples. Some of the 
most numerous taxa associated with guano deposits 
are mites (Acarina), particularly from the families 
Gamasidae, Actinedidae, Oribatidae and Armadilhdae 
(Womersley 1963a, b; Gnaspini and Trajano 2000). 
Extremely high numbers (>33 million/m 2 ) have been 
recorded on fresh guano (Harris 1973; Bellati 2001). 
Guanivores from all biogeographic regions are 
taxonomically similar, usually belonging to the same 
families. Differences, however, are found among the 
predators of guanivore communities and are often 
represented by taxa from different families depending 
on the biogeographical region (Gnaspini and Trajano 
2000). 

Bat guano micro-habitat variation 

Guano environments are extremely variable, 
consisting of numerous micro-habitats when compared 
with the majority of subterranean habitats (Harris 
1970). Bat guano deposits have been found to exhibit 
variable temperature of both the ambient air above 
deposits and within deposits (Harris 1970). In addition, 
the relative humidity, CO concentration, and ammonia 
concentration also change when bats occupy a cave 
due to their breathing and urine (Decu 1986). 
Variations in pH can be extreme, resulting in strong 
differentiation between fresh and old guano deposits. 
The annual cycle of bat roosting adds a temporal 
component to many guano deposits and also serves to 
alter air temperature in roosting chambers. Bat maternal 
chambers are especially variable when extremely large 
numbers of bats enter a chamber on an annual basis to 
birth young (Harris 1970). 

Large numbers of bats can raise the air 
temperature in a chamber by up to 10°C. This effect is 
most prevalent in high-domed chambers where heated 
air is trapped, but Harris (1970) also noted small 
increases in air temperature close to guano piles of up 
to 1.4°C due to heat released from guano breakdown. 
Increased air temperature of up to 12°C has also been 
noted in Cuban caves where large numbers of the leaf- 
nosed bat, Phyllonycteris poeyi Gundlach, roost (Decu 
1986). This temperature increase can act as a barrier 
for colonisation by generalist cavernicolous 
invertebrate species, but allows guanophilic and 
guanobitic populations to reach large numbers. 

Temperature within a guano pile can increase 
significantly with depth. Temperatures 5 cm below the 
surface of guano piles in Carrai Bat Cave, New South 
Wales are 1.7°C higher compared with surface 



Proc. Linn. Soc. N.S.W., 125, 2004 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



temperatures, and 15 cm below the surface 
temperatures are 3.0°C higher (Harris 1970). Surface 
guano temperatures have also been reported to increase 
by 9.3°C, and these increases in both surface and 
subsurface temperatures were attributed by Harris 
(1970) to the increase in the metabolic rate of the 
organisms inhabiting the guano pile. The initiation of 
growth and reproduction of mites in guano may be 
linked to the increase in temperature associated with 
bat occupation of a chamber (Harris 1971). 

Varying water content of guano due to 
desiccation with increasing age, results in noticeable 
micro-habitat differentiation. Fresh guano collected 
from the tops of piles in Bat Cave (U2), Naracoorte, 
South Australia, has been measured at up to 85% water 
by weight (Moulds 2003). Guano from the base of piles 
is a lighter grey colour due to desiccation and can 
contain as little as 6% water by weight (Moulds 2003). 
Guano moisture content increases with the birth of pups 
as their faecal matter is predominately liquid prior to 



being weened (approximately 6-8 weeks after birth 
for the large bent-wing bat Miniopterus schreibersii 
bassanii Cardinal and Christidis) (T. Moulds 
unpublished data). The surface of guano deposits 
commonly exhibit a patchwork appearance of dark 
moist areas and light grey drier areas. Different species 
within guano ecosystems prefer different micro- 
habitats. Richards (1971) noted the majority of 
guanophilic arthropods in Nullarbor Plain caves were 
only found in completely or partially dry guano. 

Guano shows a marked difference in pH 
between fresh and old deposits. Fresh guano is 
commonly basic, with the pH varying according to 
the volume of urine deposited with faeces. Fresh guano 
commonly has a pH of 8.5-9.0 that rapidly becomes 
acidic (5.0-5.5) with age and depth, although the centre 
of guano piles has a stable pH of around 4 (Harris 
1971). In bat maternity caves the pH of piles will 
gradually decrease over winter as no fresh guano is 
deposited. Data from Bat Cave (U2) (Naracoorte, 




Protochelifer cavemarum ^~% 
(Pseudoscorpionida) V/ 




Shawella douglasi 
(Blattodea) 




Ptinus exulans 
(Coleoptera) 



tf 




Uroobovella coprophila r^-$ 
(Acarina) v/ 



Figure 1. Different distribution patterns of guano associated species across Australia. 



Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Table 1. Possibly endemic, guano dependent species in Australia. 



State 


Order 


Genus and Species 


Depend 
Cave 


lance 
Guano 


Cave 


QLD 


Pseudoscorpionida 


Sathrochthonius webbi 


Tb 


Gp 


Holy Jump Lava Cave (BM1) 


QLD 


Coleoptera 


Choleva australis 


Tp 


Gp 


Royal Arch Cave (CH9) 


QLD 


Coleoptera 


Dermestes uter 


Tp 


Gp 


Royal Arch Cave (CH9) 


QLD 


Coleoptera 


Alphitobius diaperinus 


Tp? 


Gp? 


Bat Cleft (E6) 


QLD 


Coleoptera 


Omorgus costatus 


Tp 


Gp? 


Johannsens Cave (J 1-2) 


QLD 


Coleoptera 


Anomotarus subterraneus 


Tp 


Gp 


Riverton Main Cave (RN1) 


NSW 


Pseudoscorpionida 


Oratemnus cavemicola 


Tp 


Gp? 


Jump Up Cave, Gray Range 


NSW 


Pseudoscorpionida 


Sundochernes guanophilus 


Tp2 


Gb 


Fig Tree Cave (W148) 


NSW 


Pseudoscorpionida Tyrannochthonius cavicola 


Tp2 


Gb 


Grill Cave (B44) 


NSW 


Acarina 


Neotrombidium gracilipes 


Tp2 


Gb 


Fig Tree Cave (W148) 


NSW 


Acarina 


Hypoaspis annectans 


Tp 


Gp 


Carrai Bat Cave (SC5) 


Nullarbor 


Pseudoscorpionida 


Cryptocheiridium australicum 


Tp2 


Gp 


Murra-El-Elevyn Cave (N47) 


Nullarbor 


Isopoda 


Abedaioscia troglodytes 


Tb 


Gp? 


Pannikin Plain Cave (N49) 


Nullarbor 


Coleoptera 


Quedius luridipennis 


Tp? 


Gp 


Abrakurrie Cave (N3) 


VIC 


Pseudoscorpionida 


Pseudotyrannochthonius 
hamiltonsmithi 


Tp2 


Gp 


Mount Widderin Cave (HI) 


VIC 


Coleoptera 


Achosia lanigera 


Tp? 


Gp 


Wilson Cave (EB4) 


SA 


Pseudoscorpionida 


Austrochthonicus cavicola 


Tp2 


Gp 


Cathedral Cave (U12) 


SA 


Pseudoscorpionida 


Protochelifer naracoortensis 


Tp2 


Gp 


Bat Cave (U2) 


WA 


Blattodea 


Paratemnopteryx atra 


Tb 


Gp 


Mines nr Marble Bar 


Christmas 1 


'. Coleoptera 


Alphitobius laevigatus 


Unknown 


Gp 


Upper Daniel Roux Cave (CI56) 



South Australia), show that late in spring, before guano 
deposition recommences, tops of guano piles can 
become acidic, occasionally as low as pH 5.0 (Moulds 
2003). The ever changing pH of guano piles due to 
age and urine content creates marked micro-habitats 
used by differing species. 

Micro-habitat variation of bat chambers is 
further complicated by the movement of bat roosts in 
a chamber within a breeding season. These movements 
are a response to avoiding unfavourable conditions 
caused by ammonia concentrations and high local 
temperatures (Poulson 1972). 



DISTRIBUTION, BIOGEOGRAPHY AND 
ENDEMISM 

This is the first checklist for Australian guano- 
associated invertebrates. The full geographic range of 
many guanobitic and guanophilic species can now 
easily be appreciated. Many species have been shown 
to have unexpectedly wide distributions, sometimes 
spanning several climatic regions. Several possible 
explanations exist for these patterns. The lack of 
systematic searching and collation of published 
records, and collections has resulted in a poor 



understanding of many species distribution and degree 
of endemism. This is commonly combined with a lack 
of accurate identification by taxonomic experts leading 
to the lumping of several similar species into one. 
Inadequate species definitions from groups requiring 
systematic revision will also result in species being 
artificially lumped or split (eg Diptera: Phoridae, David 
McAlpine, pers. comm. 2002). A lack of collections 
from most karst areas, both above and below ground, 
is the greatest problem, resulting in large gaps in 
distributions and a poor knowledge of variation within 
species. The paucity of records among some taxa also 
provides a focal point for future collecting priorities. 
The collation of this checklist has revealed 
associations of species across wide geographic regions. 
Figure la shows the extensive range of Protochelifer 
cavernarum Beier (Pseudoscorpionida) from Jurien 
Bay, Western Australia, across southern Australia and 
north to Undara Lava Tubes in northern Queensland. 
The distribution of Shawella douglasi Princis 
(Blattodea: Blattellidae) (Fig. lb) is disjunct with 
records from northern New South Wales and Jurien 
Bay, Western Australia. This may be the result of 
misidentification, poor taxonomic description or a 
paucity of collecting between these localities, 
especially throughout northern Australia. Despite a 



Proc. Linn. Soc. N.S.W., 125, 2004 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



number of invertebrate collections from the Nullarbor 
karst no individuals have been recorded, possibly due 
to extremely small populations of troglobitic species 
and the extremely large size of the karst area concerned. 
Several species including Ptinus exulans Erichson 
(Coleoptera: Anobiidae) show very wide distributions 
from mid-north New South Wales across the Nullarbor 
Plain to the west coast of Western Australia (Fig. lc). 
The distribution of U. coprophila (Acarina: 
Urodinychidae) (Fig. Id) is directly linked to the 
distribution of maternal sites for the large bent-wing 
bat M. schreibersii. The single record of this species 
from Undara (north Queensland) may be spurious, a 
misidentification or an individual transported via 
phoresy, especially as no records exist between 
southern and northern Queensland despite large bat 
maternity caves around Rockhampton. These data raise 
further questions regarding the colonisation of guano 
deposits by invertebrates and the boundaries of 
possibly ill-defined species concepts. 

Endemic status of guano species, has, in the 
past been assigned without a full understanding of the 
distribution of Australian guano fauna. This is apparent 
for the maternal chamber of Bat Cave (U2), 
Naracoorte, where previous studies (Hamilton-Smith 
2000), identified 'several endemic species' to the 
maternal chamber or Bat Cave as a whole. This 
checklist has shown that Bat Cave contains only a 
single endemic species, Protochelifer naracoortensis 
Beier, and this pseudoscorpion may possibly be found 
in other caves in the continuous karst of the Otway 
Basin. Bat Cave does, however, form the most diverse 
guanophilic arthropod community in Australia. This 
highlights the amount of assumed knowledge 
concerning guano invertebrates in Australia and their 
distribution. The number of endemic species to specific 
bat caves is currently unknown but is almost certainly 
significantly lower than previously believed. Several 
species have been identified as possessing restricted 
distributions and guano dependence, although none 
can yet be positively identified as endemic (Table 1). 
The restricted distribution status of all species listed 
in Table 1 is tentative and more extensive collecting, 
both above and below ground, must be undertaken 
before distribution can be confirmed. This is especially 
true for troglophilic species as epigean occurrence of 
these species will effect their endemic status. The 
degree of a species' guano dependence will also affect 
its endemic status and more ecological knowledge is 
required to confirm species habits. Species confined 
to single caves or isolated areas are more likely to be 
endemic when combined with guano dependence. 
Only Fig Tree Cave (W148) (Wombeyan, NSW) and 
Royal Arch Cave (CH9) (Chillagoe, QLD) are found 



to contain two species showing both restricted 
distribution and guano dependence (Table 1). 

The presence of nematodes is almost a 
certainty in guano caves as they are almost ubiquitous 
in every other habitat both above and below ground. 
Despite this the records of nematodes from guano are 
extremely limited primarily because the majority of 
caves and karst areas remain completely unsampled 
for these invertebrates. Nematodes play a potentially 
important role in the micro-habitat of guano piles and 
have been recorded in large numbers from overseas 
caves (Decu 1986). Nematodes are also believed to be 
one of the first colonisers of new bat caves, being 
deposited by in urine and faeces (Decu 1986). Further 
sampling of Australian cave guano will almost 
certainly reveal a greater diversity of species. Currently 
no free living nematodes have been recorded by the 
author from Bat Cave, Naracoorte despite several 
collection events. 

Currently no guano invertebrates are recorded 
from Tasmania, primarily due to the absence of cave- 
dwelling bats. The possibility remains however, that 
guano communities occur in orthopteran guano or 
other invertebrate guano deposits or even bird guano. 
The guanophilic mite Macrocheles tenuirostris Krantz 
and Filipponi was first recorded from mutton bird nests 
in Tasmania and has since been collected from bat 
guano in Victorian and New South Wales caves. 
Further field observations within Tasmanian caves may 
yet reveal these communities. 

Opportunities for future research in this field 
are vast with only limited knowledge existing for most 
karst areas. The ecological classification for many 
species is poorly known and this will only be achieved 
through increased observations in situ. The 
microbiology of guano deposits also remain very 
poorly known in Australian, as well as in overseas 
caves. Many karst areas remain completely unstudied 
biologically, especially with regard to the diversity of 
invertebrate guano communities. 



SYSTEMATIC CHECK LIST OF AUSTRALIAN 
GUANO INVERTEBRATES 

This checklist includes all Australian 
cavernicolous species found in association with guano 
from both caves and mines. Records have been 
compiled from the speleological literature (both 
scientific and amateur), unpublished records, and 
personal observations. Parasites of cave-dwelling 
mammals (bats) have been included as they are often 
found in guano, although their potential roles in guano 
ecosystems is currently unknown. Taxa are arranged 



Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



systematically by Phylum, Class and Order then 
alphabetically by Family. Undetermined taxa have 
been placed at the end of their respective order or 
family. Due to changes in taxonomy and higher 
systematics of many taxa the names and position of 
species can be uncertain. This checklist has adopted 
the most recent higher classifications attainable and 
many old names have been updated to reflect changes 
in the literature. Many groups in this checklist are in 
need of revision and so some species concepts may be 
altered in the future resulting in the splitting of some 
species and the lumping of others. This will obviously 
affect the distribution of species as presented in this 
work. 

Cave names and numbers following the 
Australian Karst Index (Mathews 1985), and are listed 
for all species' records along with appropriate 
references. Records from caves in the Nullarbor Plain, 
southern Australia, have not been divided along state 
boundaries in order to reflect the extremely large and 
continuous nature of this karst area. Taxa previously 
considered to be obvious accidentals to cave 
environments have been excluded from this checklist. 

The following ecological classification is 
modified from Hamilton-Smith (1967), and Gnaspini 
and Trajano (2000), and is based on the degree of cave 
and guano dependence of taxa. Abbreviations are those 
used in the checklist. 

Trogloxene (Tx): an organism that regularly uses the 
cave environment for part of its lifecycle or as shelter 
but must leave the cave to feed and or breed. 
1 st order Troglophile (Tpl): an organism that can 
complete its entire lifecycle within a cave but possess 



no specific adaptations to the cave environment and 

recorded in both epigean and hypogean habitats. 

2 nd order Troglophile (Tp2): an organism that can 

complete its entire lifecycle within a cave but possess 

no specific adaptations to the cave environment and 

recorded only from hypogean habitats. 

Troglobite (Tb): obligate cavernicolous organisms 

that possess specific adaptations to the cave 

environment. 

Guanoxene (Gx): an organism that may use guano 

for reproduction and/or feeding but requires other 

substrates to complete its life cycle. 

Guanophile (Gp): an organism that inhabits and 

reproduces both in guano piles as well as other 

substrates within a cave. 

Guanobite (Gb): an organism that requires guano 

deposits to complete its entire life cycle. 

Bat Parasite (P): an animal that is an obligate bat 

parasite requiring bats to complete its lifecycle. 

Ecological classifications have been assigned 
to taxa wherever possible. These designations were 
made using available knowledge concerning 
behaviour, life history, and distribution within caves. 
However, information regarding species' ecology was 
found to be lacking or minimal in most cases. Because 
of such constraints some taxa have not been assigned 
a guano classification. Further, information on other 
taxa was insufficient to confirm their association with 
guano ecosystems. Thus, taxa previously recorded only 
from guano caves, but without a confirmed association 
with guano, have been included for completeness even 
though some of these species may be unassociated with 
guano. 



Phylum Platyhelminthes 

Class Tubellaria 
Order undetermined 

Undetermined genus and species, Tx, Gx?. VICTORIA: Dickson Cave (M30), Murrindal (Yen and 
Milledge 1990). 

Phylum Nemathelminthes 

Class Nematoda 
Order Strongyloidea 

Trichostrongylidae 

Nycteridostrongylus unicollis Baylis, Tx, Gx, P. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Hamilton-Smith unpublished data). 

Molinostrongylus dollfusi Mawson, Tx, Gx, P. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Hamilton-Smith unpublished data). 



Proc. Linn. Soc. N.S.W., 125, 2004 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Order Undetermined 

?Rhabditida 

Undetermined genus and species, Gp?. VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds 
unpublished data), bacterial feeder (K. Davies pers. comm. 2003). 

Undetermined Family 

Undetermined genus and species, Gp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard 
Creek (Harris 1970). 

Undetermined genus and species, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 
1998). 

Phylum Mollusca 

Class Gastropoda 
Order Stylommatophora 

Charopidae 

Elsotherafunera Cox, Gx?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith 
unpublished data); VICTORIA: Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades 
of Death Cave (M3), Murrindal (Yen and Milledge 1990); Anticline Cave (Mil), Murrindal (Yen 
and Milledge 1990). 

Undetermined Family 

Undetermined genus and species, Tx, Gx?. NORTHERN TERRITORY: Cutta Cutta Cave (Kl), 
Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Cam Dum (E15), Mount Etna 
(Hamilton-Smith unpublished data). 

Phylum Annelida 

Class Oligochaeta 
Order Haplotaxida 

Lumbricidae 

Undetermined genus and species, Tp, Gx?. VICTORIA: Wilson Cave (EB4), East Buchan (Yen and 
Milledge 1990). 

Order Undetermined 

Undetermined genus and species, Tp?, Gx?. QUEENSLAND: Four Mile Cave (C14), Camooweal 
(Hamilton-Smith unpublished data). 

Phylum Arthropoda 

Class Arachnida 
Order Scorpionida 

Undetermined Family 

Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (Mil), Murrindal (Yen and 
Milledge 1990). 

Order Araneae 

Agelenidae 

Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (Mil), Murrindal (Yen and 
Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). 

Amaurobiidae 

Undetermined genus and species, Gx?. VICTORIA: Spring Creek Cave (Bl), Buchan (Yen and 



Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 

Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), 
East Buchan (Yen and Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990). 

Ctenizidae 

Misgolas sp., NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Gray 1973b). 

Cyatholipidae 

Undetermined genus and species, Gx?. VICTORIA: Lilly Pilly Cave (M8), Murrindal (Yen and 
Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), 
Murrindal (Yen and Milledge 1990). 

Cycloctenidae 

Cyclotenus abyssinus Urquhart, Tp. VICTORIA: Shades of Death Cave (M3), Murrindal (Hamilton- 
Smith unpublished data). 

Toxopsioides sp., Tp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Gray 
1973b); Yessabah Bat Cave (YE1), Yessabah (Gray 1973b). 

Undetermined genus and species, Gx?. VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 
1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave (M3), 
Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); 
Dickson Cave (M30), Murrindal (Yen and Milledge 1990). 

Desidae 

Badumna socialis Rainbow, Tp, Gx?. NEW SOUTH WALES: Chalk Cave (B26), Bungonia 
(Hamilton-Smith unpublished data). 

Colcarteria carrai Gray, Tp?. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(Gray 1992). 

Colcarteria yessabah Gray, Tp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(Gray 1992). 

Dictynidae 

Undescribed genus and species, Gx?. VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 
1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990). 

Filistatidae 

Undescribed genus and species, Tp2. WESTERN AUSTRALIA: Cape Range peninsula (Gray 
1994). 

Gradungulidae 

Progradungula carraiensis Forster and Gray, Tpl, Gp . NEW SOUTH WALES: Carrai Bat Cave 
(SC5), Stockyard Creek (Forster et al. 1987). 

Linyphiidae 

Laetesia weburdi Urquhart, Gx?. NEW SOUTH WALES: Jenolan Caves (Hamilton-Smith 
unpublished data). 

Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (Mil), Murrindal (Yen and 
Milledge 1990). 

Lycosidae 

Lycosa speciosa Koch, Tpl. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(Gray 1973b). 

Proc. Linn. Soc. N.S.W., 125, 2004 9 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Mimetidae 

Australomimetus maculosus Rainbow, Tp. NEW SOUTH WALES: Yessabah Bat Cave (YE1), 
Yessabah (Gray 1973b); Colong Main Cave (CGI), Colong (Hamilton-Smith unpublished data); 
Jenolan Caves (Hamilton-Smith unpublished data). 

Undetermined genus and species, Gx?. VICTORIA: Spring Creek Cave (Bl), Buchan (Yen and 
Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990). 

Pholcidae 

Physocyclus sp., NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Gray 1973b); 
Colong Main Cave (CG3), Colong (Gray 1973b). 

Psilochorus sp., NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Gray 1973b) . 

Pisauridae 

Undetermined genus and species, NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Gray 
1973b); Carrai Bat Cave (SC5), Stockyard Creek (Gray 1973b). 

Salticidae 

Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (Mil), Murrindal (Yen and 
Milledge 1990). 

Segestriidae 

Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (Mil), Murrindal (Yen and 
Milledge 1990). 

Stiphidiidae 

Stiphidon sp., Gx?. NEW SOUTH WALES: Colong Cave (CGI), Colong (Hamilton-Smith 
unpublished data). 

Theridiidae 

Theridon sp., Tp, Gp. NEW SOUTH WALES: Colong Cave (CGI), Colong (Hamilton-Smith 
unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Steatoda sp., Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Theridiosomatinae 

Undetermined genus and species, Gp?. NEW SOUTH WALES: Colong Cave (CGI), Colong 
(Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished 
data). 

Uloboridae 

Philoponella patherinus Keyserling, Tp. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data). 

Undetermined Family 

Undetermined genus and species, Gp?. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- 
Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); 
Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Cave (CGI), Colong 
(Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton-Smith unpublished 
data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data); 
Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave 
(Tl), Tuglow (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton- 



10 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 

Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith 
unpublished data); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data); Fig Tree 
Cave (W148), Wombeyan (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Cutta 
Cutta Cave (Kl), Katherine (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Abrakurrie 
Cave (N3) (Hamilton-Smith unpublished data); Madura Cave (Madura 6 Mile Cave) (N62) 
(Hamilton-Smith unpublished data); QUEENSLAND: Four Mile Cave (C14), Camooweal 
(Hamilton-Smith unpublished data); Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith 
unpublished data); Holy Jump Lava Cave (BM1), Bauer's Mountain (Hamilton-Smith unpublished 
data); Barker's Cave (U34), Undara (Hamilton-Smith unpublished data); Johannsen's Cave (Jl-2), 
Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), 
Mt Etna (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith 
unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Piglet 
Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data); Ilium Cave (E31), Mount 
Etna (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith 
unpublished data); SOUTH AUSTRALIA: Snowflake Cave (LI), Glenelg River (Hamilton-Smith 
unpublished data); Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan 
(Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Trogdip 
Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Shades of Death Cave (M3), 
Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); 
Anticline Cave (Mil), Murrindal (Yen and Milledge 1990); SSS Cave (M44), Murrindal 
(Hamilton-Smith unpublished data); Nargun's Cave (NN1), Nowa Nowa (Hamilton-Smith 
unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Mt Widderin Cave 
(HI), Skipton (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton- 
Smith unpublished data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); 
Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). 

Order Opilionida 

Triaenoncychidae 

Holonuncia cavernicola Forster, Tp2. NEW SOUTH WALES: Basin Cave (W4), Wombeyan 
(Hamilton-Smith 1967); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data). 

Holonuncia seriata Roewer, Tpl, Gx. NEW SOUTH WALES: Bungonia various caves (Eberhard 
1998). 

Undetermined genus and species, Tp, Gp. VICTORIA: Moon Cave (B2), Buchan (Yen and 
Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave 
(M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 
1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal 
(Yen and Milledge 1990). 

Undetermined Family 

Undetermined genus and species, Tp, Gp? NEW SOUTH WALES: The Drum Cave (B13), 
Bungonia (Hamilton-Smith unpublished data); Chalk Cave (B26), Bungonia (Hamilton-Smith 
unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Cliefden Main 
Cave (CL1), Chef den (Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton- 
Smith unpublished data); Colong Main Cave (CGI), Colong (Hamilton-Smith unpublished data); 
Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data); 
Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith unpublished data); Glen 
Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (Tl), 
Tuglow (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith 
unpublished data); Yessabah Bat Cave (YE1), Yessabah (Hamilton-Smith unpublished data); 
NULLARBOR PLAIN: Lynch Cave (N60) (Hamilton-Smith unpublished data); QUEENSLAND: 
Johannsen's Cave (Jl-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); 



Proc. Linn. Soc. N.S.W., 125, 2004 1 1 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Unnamed 
Cave (NG1), New Guinea Ridge (Hamilton-Smith unpublished data). 

Order Pseudoscorpionida 

Atemnidae 

Oratemnus cavernicola Beier, Tp, Gp?. NEW SOUTH WALES: Jump Up Cave, Gray Range (Beier 
1976). 

Cheiridiidae 

Cryptocheiridium australicum Beier, Tp2, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave 
(N47) (Richards 1971). 

Cheliferidae 

Protochelifer naracoortensis Beier, Tp2, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Beier 1968; Bellati et al. 2003). 

Protochelifer cavernarum Beier, Tp2, Gb. NEW SOUTH WALES: Murder Cave (CL2), Cliefden 
(Beier 1967, 1968); Island Cave (CL6), Cliefden (Hamilton-Smith unpublished data); Belfry Cave 
(TR2), Timor (Beier 1967); Ashford Caves, Ashford (Beier 1968); NULLARBOR PLAIN: Warbla 
Cave (Nl) (Richards 1971); Weebuddie [Weebubbie, sic] Cave (N2) (Beier 1975); Abrakurrie Cave 
(N3) (Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); Mullamullang Cave (N37) 
(Richards 1971); Lynch Cave (N60) (Richards 1971); QUEENSLAND: Taylor Cave (4U4), Undara 
(Howarth 1988); Collins Cave No.l, Undara (Howarth 1988); VICTORIA: Clogg's Cave (EB2), 
East Buchan (Beier 1968); WESTERN AUSTRALIA: Gooseberry Cave (Jl), Jurien Bay (Beier 
1968); Eneabba Caves (El-3), Eneabba (Lowry 1996); Arramall Cave (E22), Eneabba (Lowry 
1996); River Cave (E23), Eneabba (Lowry 1996); Weelawadji Cave (E24) Eneabba (Lowry 1996); 
Super Cave (SHI), South Hill River (Hamilton-Smith unpublished data). 

Protochelifer sp. Tp, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Bellati et al. 
2003). 

Chernetidae 

Sundochernes guanophilus Beier, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Beier 1967). 

Troglochernes imitans Beier, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elvyn Cave (N47) (Beier 
1975); Cocklebiddy Cave (N48) (Beier 1975); Pannikin Plain Cave (N49) (Beier 1975); Dingo 
Cave (Dingo-Donga) (N160) (Richards 1971). 

Chthoniidae 

Austrochthonius cavicola Beier, Tp, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte 
(Beier 1968). 

Paraliochthonius cavicolus Beier, Tp2, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998). 

Pseudotyrannochthonius hamiltonsmithi Beier, Tp2, Gp. VICTORIA: Mt Widderin Cave (HI), 
Skipton (Beier 1968). 

Sathrochthonius tuena Chamberlin, Tp2, Gp. NEW SOUTH WALES: Basin Cave (W4), 
Wombeyan (Beier 1967), Deua Cave (DEI), Deua (Eberhard and Spate 1995); Punchbowl Cave 
(WJ8), Wee Jasper (Beier 1968); Imperial Cave (J4), Jenolan (Hamilton-Smith 1967; Gibian et al. 
1988); Southern Limestone, Jenolan (Hamilton-Smith 1967; Beier 1968; Gibian et al. 1988); 
Paradox Cave (J48), Jenolan (Hamilton-Smith unpublished data). 



12 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Sathrochthonius webbi Muchmore, Tb, Gp. QUEENSLAND: Holy Jump Lava Cave (BM1), 
Bauer's Mountain southern Queensland (Muchmore 1982). 

Tyrannochthonius cavicola Beier, Tp2, Gb. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Beier 1967; Harvey 1989). 

Undetermined Family 

Undetermined genus and species, Tp, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Royal Arch Tower, 
Chillagoe (Matts 1987). 

Undetermined genus and species, Tp?, Gx. VICTORIA: Anticline Cave (Mil), Murrindal (Yen and 
Milledge 1990). 

Mites 

The mites have been arranged according to the higher classification used by Halliday (1998). Many changes 
to nomenclature have occurred since previous checklists of cavernicolous fauna have been published so the 
family placement of some species has been updated to reflect this. Previous family placements have not been 
recorded but where synonymy has occured the old name (either family or genus) has been included in 
brackets. Previous generic placements have been recorded in brackets with the prefix "=". 

Order Acariformes 
Suborder Astigmata 

Histiostomatidae 

Histiostoma sp. NULLARBOR PLAIN: Mullamullang Cave (N37) (Hamilton-Smith unpublished 
data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Rosensteiniidae 

Nycteriglyphus (Coproglyphus) dewae Zakhvatkin, Tp2, Gb. NEW SOUTH WALES: Basin Cave 
(W4), Wombeyan (Womersley 1963a; Richards 1967b); Fig Tree Cave (W148), Wombeyan 
(Womersley 1963a; Richards 1967b); Railway tunnel, North Sydney (Womersley 1963a); SOUTH 
AUSTRALIA: Bat Cave (U2), Naracoorte (Womersley 1963a). 

Nycteriglyphus sp., Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971); 
Dingo Cave (160) (Richards 1971). 

Glycyphagus sp., Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971); 
Dingo Cave (N160) (Richards 1971). 

Suborder Prostigmata 

Labidostomidae 

Undetermined genus and species. NEW SOUTH WALES: Island Cave (CL6), Cliefden (Hamilton- 
Smith unpublished data). 

Neotrombidiidae 

Neotrombidium gracilare Womersley, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Womersley 1963a); Murder Cave (CL2), Cliefden (Womersley 1963a); Punchbowl 
Cave (WJ8), Wee Jasper (Womersley 1963a); VICTORIA: O'Rourke's Cave (B12), Buchan 
(Hamilton-Smith 1967); Wilson Cave (EB4), East Buchan (Hamilton-Smith 1967). 

Neotrombidium gracilipes Womersley, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Hamilton-Smith 1967). 



Proc. Linn. Soc. N.S.W., 125, 2004 13 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

Neotrombidium neptunium Southcott, VICTORIA: Clogg's Cave (EB2), East Buchan (Hamilton- 
Smith unpublished data). 

Neotrombidium sp., Tp, Gb. NULLARBOR PLAIN: Firestick Cave (N70) (Richards 1971); Dingo 
Cave (N160) (Richards 1971). 

Trombiculidae 

Rudnicula barbarae Domrow (= Trombicula), Tx, Gx, P. NORTHERN TERRITORY: Kuhinoor 
Mine, Pine Creek (Hamilton-Smith unpublished data). 

Trombicula thomsoni Womersley, Tx, Gx, P. NEW SOUTH WALES: Bonalbo Colliery (Hamilton- 
Smith unpublished data); Riverton (Hamilton-Smith unpublished data); NORTHERN 
TERRITORY: Kuhinoor Mine, Pine Creek (Hamilton-Smith unpublished data). 

Trombicula dewae Domrow, Tx, Gx, P. NORTHERN TERRITORY: Kuhinoor Mine, Pine Creek 
(Hamilton-Smith unpublished data). 

Order Parasitiformes 
Suborder Ixodida 

Argasidae 

Argas sp., Tx, Gx, P. QUEENSLAND: Johannsen's Cave (Jl-2), Limestone Ridge, Rockhampton 
(Hamilton-Smith unpublished data). 

Ixodidae 

Amblyomma moreliae Koch, Gx, P. QUEENSLAND: Johannsen's Cave (Jl-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data). 

Ixodes simplex simplex Neumann, Gx, P. Bat parasite in eastern Australia (Hamilton-Smith 1966b; 
Eberhard 1998); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished 
data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Spring 
Creek Cave (Bl), Buchan (Hamilton-Smith unpublished data); Slocombe's Cave (BA1), The Basin 
(Hamilton-Smith unpublished data); Anticline Cave (Mil), Murrindal (Hamilton-Smith 
unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data); Starlight 
Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Grassmere Cave (W6), 
Warrnambool (Hamilton-Smith unpublished data). 

Undetermined genus and species, Gx, P. QUEENSLAND: Clam Cavern (CH26), Walkunder 
Tower, Chillagoe (Matts 1987); Spatial Cavern (CH41), Walkunder Tower, Chillagoe (Marts 1987); 
Royal Arch Cave (CH9), Royal Arch Tower, Chillagoe (Matts 1987); VICTORIA: Nargun's Cave 
(NN1), Nowa Nowa (Hamilton-Smith unpublished data) 

Suborder Mesostigmata 

Ameroseiidae 

Ameroseius plumosus Oudemans, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et 
al. 2003). 

Laelapidae 

Cosmolaelaps sp., Tp2, Gb. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton- 
Smith 1967); QUEENSLAND: Railway tunnel, Samford (Hamilton-Smith 1967). 

Hypoaspis (Gaeolaelaps) annectans Womersley, Tp, Gp. NEW SOUTH WALES: Carrai Bat Cave 
(SC5), Stockyard Creek (Harris 1971). 

Hypoaspis (Gaeolaelaps) sp.l, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 

14 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Hypoaspis (Gaeolaelaps) sp.2, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 

Hypoaspis (Gaeolaelaps) sp., Tp2, Gb. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- 
Smith 1967). 

Ichoronyssus (Pleisiolaelaps) miniopteri (Zumpt and Patterson 1952) (= Neospinolaelaps, 
Spinolaelaps), Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998); 
Bonalbo Colliery (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), 
Naracoorte (Hamilton-Smith unpublished data). 

Ichoronyssus (Pleisiolaelaps) aristippe Domrow, NEW SOUTH WALES: Cheitmore Cave, 
Cheitmore (Hamilton-Smith unpublished data); Wombeyan Caves (Hamilton-Smith unpublished 
data); Bonalbo Colliery (Hamilton-Smith unpublished data). 

Macrochelidae 

Macwcheles spatei Halliday, Tpl, Gp. NEW SOUTH WALES: Deua Cave (DEI), Deua National 
Park (Halliday 2000). 

Macrocheles tenuirostris Krantz and Filipponi, Tpl, Gp. NEW SOUTH WALES: Paradox Cave 
(J48), Jenolan Caves (Halliday 2000); Cleatmore Cave, Deua National Park (Halliday 2000); 
Colong Cave, Woofs Cavern (CGI), Colong (Halliday 2000); Church Cave (WJ31), Wee Jasper 
(Halliday 2000); TASMANIA: Fisher Island, in nests and burrows of muttonbird (Krantz and 
Fillipponi 1964); VICTORIA: Panmure Cave (H5), Warrnambool (Hamilton-Smith 1967). 

Macronyssidae 

Macronyssus aristippe Domrow (= Ichoronyssus), Tp, Gx, P. NEW SOUTH WALES: Bungonia 
various caves (Eberhard 1998). 

Trichonyssus australicus Womersley, Tx, Gx, P. NULLARBOR PLAIN: Warbla Cave (Nl) 
(Hamilton-Smith unpublished data). 

Parantennulidae 

Micromegistus gourlayi Womersley. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 

Parasitidae 

7Eugamasus sp., Tp, Gp. NULLARBOR PLAIN: Dingo Cave (N160) (Richards 1971). 

Sejidae (Ichthyostomatogastridae) 

Asternolaelaps australis Womersley and Domrow, Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2) 
Naracoorte (Womersley and Domrow 1959; Hamilton-Smith 1967); VICTORIA: O'Rourkes Cave 
(B12), Buchan (Hamilton-Smith 1967). 

Spinturnicidae 

Spinturnix psi Kolenati, Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Eberhard 
1998). 

Undetermined genus and species, Tp, Gx, P. NEW SOUTH WALES: Colong Main Cave (CGI), 
Colong (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie Cave (N2) 
(Hamilton-Smith unpublished data); Murra-El-Elevyn Cave (N47) (Hamilton-Smith unpublished 
data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); 
Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton (Hamilton-Smith 



Proc. Linn. Soc. N.S.W., 125, 2004 15 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 
unpublished data); VICTORIA: Spring Creek Cave (Bl), Buchan (Hamilton-Smith unpublished 
data); WESTERN AUSTRALIA: Stockyard Cave (E3), Eneabba (Hamilton-Smith unpublished 
data). 

Urodinychidae 

Uroobovella (Austruropoda) coprophila Womersley (= Cilliba), Tp2, Gp. NEW SOUTH WALES: 
Cave C4, Comboyne (Smith 1982b); Carrai Bat Cave (SC5), Stockyard Creek (Harris 1973); 
Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Church Cave (WJ31), 
Wee Jasper (Hamilton-Smith 1966b, 1967); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith 
1966b, 1967); Cheitmore Cave, Cheitmore (Hamilton-Smith unpublished data); QUEENSLAND: 
Arch Cave (U22), Undara (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), 
Riverton (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Bellati et al. 2003); VICTORIA: Anticline Cave (Mil), Murrindal (Hamilton-Smith 1967). 

Genus and species undetermined, NEW SOUTH WALES: Deua Cave (DEI), Deua (Eberhard and 
Spate 1995). 

Undetermined Family 

Undetermined sp. 1, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 
1971). 

Undetermined sp. 2, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 
1971). 

Undetermined Acarina 

Undetermined Family 

Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Grimes Cave (CI53) 
(Humphreys and Eberhard 2001). 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Gable Cave (CL7), Cliefden 
(Hamilton-Smith unpublished data); NORTHERN TERRITORY: Kintore Cave (K2), Katherine 
(Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton- 
Smith unpublished data); Murra-El-Elevyn Cave (N47) (Hamilton-Smith unpublished data); 
QUEENSLAND: Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Asbestos mine near Arkaba, Flinders 
Ranges (Hamilton-Smith unpublished data); Drop Drop Cave (L29), Lower south east (Hamilton- 
Smith unpublished data); Joanna Bat Cave (U38), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Spring Creek Cave (Bl), Buchan (Yen and Milledge 1990); O'Rourkes Cave (B12), 
Buchan (Hamilton-Smith unpublished data); Mabel Cave (EB1), East Buchan (Yen and Milledge 
1990); Wilson's Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip Cave 
(EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), Murrindal (Yen 
and Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990); Dickson Cave 
(M30), Murrindal (Yen and Milledge 1990); Nargun's Cave (NN1), Nowa Nowa (Hamilton-Smith 
unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Grassmere Cave 
(W6), Warrnambool (Hamilton-Smith unpublished data). 

Class Crustacea 
Order Isopoda 

Armadillidae 

Merulana sp. nov., Tp. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Dennis 1986). 

Oniscidae 

Plymophiloscia sp. Vandel, Tp, Gp. NULLARBOR PLAFN: Pannikin Plain Cave (N49) (Richards 
1971; Gray 1973a). 

16 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Undetermined genus and species, Tp, Gp. QUEENSLAND: Johannsen's Cave (Jl-2), Limestone 
Ridge, Rockhampton (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna 
(Hamilton-Smith unpublished data); Cam Dum (E15), Mount Etna (Hamilton-Smith unpublished 
data); VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data). 

Philosciidae 

Abebaioscia troglodytes Vandel, Tb, Gp?. NULLARBOR PLAIN: Pannikin Plain Cave (N49) 
(Vandel 1973). 

Eurygastor montanus troglophilus Vandel, Tp?. VICTORIA: Anticline Cave (Mil), Murrindal 
(Vandel 1973). 

Laevophiloscia dongarrensis Wahrberg, Tp, Gx?. WESTERN AUSTRALIA: Yanchep Cave 
(YN16), Yanchep (Vandel 1973); Minnie's Grotto (YN28), Yanchep (Vandel 1973); Gooseberry 
Cave (Jl), Jurien Bay (Vandel 1973). 

Laevophiloscia hamiltoni Vandel, Tp, Gx. WESTERN AUSTRALIA: Weelawadji Cave (E24), 
Eneabba (Vandel 1973); Labyrinth Cave (AU16), Augusta (Vandel 1973) 

Laevophiloscia michaelseni Vandel, Tp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Vandel 
1973). 

Porcellionidae 

Porcellio scaber Latreille, Tpl. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 

Undetermined Family 

Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Ashford Main Cave (AS1), 
Ashford (Hamilton-Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith 
unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Cliefden Main 
Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Cave C4, Comboyne (Hamilton-Smith 
unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith 
unpublished data); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith 
unpublished data); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); 
Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave 
(Tl), Tuglow (Hamilton-Smith unpublished data); Piano Cave (Long Cave) (WA12), Walli 
(Hamilton-Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith 
unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith 
unpublished data); Yessabah Bat Cave (YE1), Yessabah (Hamilton-Smith unpublished data); 
NORTHERN TERRITORY: Cutta Cutta Cave (Kl), {Catherine (Hamilton-Smith unpublished data); 
NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith unpublished data); Cocklebiddy 
Cave (N48) (Hamilton-Smith unpublished data); QUEENSLAND: Barker's Cave (U34), Undara 
(Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished 
data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); SOUTH 
AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); Cathedral Cave 
(U12), Naracoorte (Hamilton-Smith unpublished data); Fox Cave (U22), Naracoorte (Hamilton- 
Smith unpublished data); Cave Park Cave (U37), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Spring Creek Cave (Bl), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East 
Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); 
Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Anticline Cave (Mil), Murrindal 
(Yen and Milledge 1990); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); 
WESTERN AUSTRALIA: Drovers Cave (J2), Jurien Bay (Hamilton-Smith unpublished data); 
Stockyard Cave (E3), Eneabba (Hamilton-Smith unpublished data). 



Proc. Linn. Soc. N.S.W., 125, 2004 17 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Order Amphipoda 

Undetermined Family 

Undetermined genus and species, Tp, Gx. VICTORIA: Wilson Cave (EB4), East Buchan (Yen and 
Milledge 1990). 

Class Myriapoda 
Order Diplopoda 

Undetermined Family 

Undetermined genus and species, Tp2, Gx. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998). 

Undetermined genus and species, Tp?, Gx. NEW SOUTH WALES: Island Cave (CL6), Cliefden 
(Hamilton-Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith 
unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Paradox Cave 
(J48), Jenolan (Hamilton-Smith unpublished data); Moparabah Cave (Temagog Cave) (MP1), 
Moparabah (Hamilton-Smith unpublished data); Carrai Bat Cave (SC5), Stockyard Creek 
(Hamilton-Smith unpublished data); Belfry Cave (TR2), Timor (Hamilton-Smith unpublished data); 
Tuglow Cave (Tl), Tuglow (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan 
(Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith 
unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Willi 
Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); Yessabah Bat 
Cave (YE1), Yessabah (Hamilton-Smith unpublished data); QUEENSLAND: Barker's Cave (U34), 
Undara (Hamilton-Smith unpublished data); Johannsen's Cave (Jl-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna 
(Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished 
data); Piglet Help! Help! Cave (El 7), Mount Etna (Hamilton-Smith unpublished data); Jolly Roger 
Cave (E29), Mountt Etna (Hamilton-Smith unpublished data); Glen Lyon River Cave (GL1), Glen 
Lyon (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith 
unpublished data); VICTORIA: Spring Creek Cave (Bl), Buchan (Yen and Milledge 1990); Mabel 
Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and 
Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990); Nargun's Cave (NN1), 
Nowa Nowa (Hamilton-Smith unpublished data). 

Order Chilopoda 

S colopendromorpha 

Undetermined genus and species. NULLARBOR PLAIN: Mullamullang Cave (N37) (Richards 
1971). 

Undetermined Family 

Undetermined genus and species, Gp?. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- 
Smith unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith 
unpublished data); Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); 
Moparabah Cave (MP1), Moparabah (Hamilton-Smith unpublished data); Belfry Cave (TR2), 
Timor (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Cutta Cutta Cave (Kl), 
Katherine (Hamilton-Smith unpublished data); Kintore Cave (K2), Katherine (Hamilton-Smith 
unpublished data); NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Hamilton-Smith unpublished 
data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); 
Johannsen's Cave (Jl-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 



Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Superclass Hexapoda 
Class Insecta 

Order Collembola 

Armadillidae 

Buddelundia albomarginata Wahrberg, Tp, Gx?. NULLARBOR PLAIN: Murrawyinee [sic] No.l 
Cave (N7) (Vandel 1973); Cocklebiddy Cave (N48) (Vandel 1973); Lynch Cave (N60) (Vandel 
1973); Madura Cave (N62) (Vandel 1973); Old Homestead Cave (N83) (Vandel 1973); Unnamed 
cave (N140) (Vandel 1973). 

Entomobryidae 

Lepidocyrtus sp., Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Lepidosira australica Schott, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 

Undetermined genus and species, Gp?. NEW SOUTH WALES: Belfry Cave (TR2), Timor (James 
et al. 1976); Chalk Cave (B26), Bungonia (Hamilton-Smith unpublished data). 

Hypogastruridae 

Hypogastrura sp., NEW SOUTH WALES: Grill Cave (B44), Bungonia (Wellings 1977); SOUTH 
AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Isotomidae 

Folsomia Candida Willem, Tp. NEW SOUTH WALES: Paradox Cave (J48), Jenolan (Eberhard 
1993); Imperial Cave (J4), Jenolan (Eberhard and Spate 1995); Tuglow Main Cave (Tl), Tuglow 
(Eberhard 1993); Jillebean Cave (Y22), Yarrangobilly (Eberhard 1993). 

Paronellidae 

Undetermined genus and species, NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan 
(Eberhard and Spate 1995). 

Undetermined Family 

Undetermined genus and species, Tp, Gp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) 
(Richards 1971); Lynch Cave (N60) (Richards 1971); Dingo Cave (N160) (Richards 1971); 
VICTORIA: SSS Cave (M44), Murrindal (Hamilton-Smith unpublished data); Mt Widderin Cave 
(HI), Skipton (Hamilton-Smith unpublished data). 

Undetermined genus and species, Tp?, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); Colong Main Cave (CGI), Colong (Hamilton-Smith 
unpublished data); Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished 
data); NORTHERN TERRITORY: 16 Mile Cave, Katherine (Hamilton-Smith unpublished data); 
QUEENSLAND: Speaking Tube (E7), Mount Etna (Hamilton-Smith unpublished data); SOUTH 
AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: 
Moon Cave (B2), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and 
Milledge 1990); Wilson's Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip 
Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), Murrindal 
(Yen and Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990); Panmure 
Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 

Order Diplura 

Undetermined family 

Undetermined genus and species, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 



Proc. Linn. Soc. N.S.W., 125, 2004 19 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Order Blattodea 

Blattellidae 

Neotemnopteryx australis Saussure, Tp, Gp. NEW SOUTH WALES: Moparabah Cave (Temagog 
Cave) (MP1), Moparabah (Hamilton-Smith 1967); Cave C4, Comboyne (Hamilton-Smith 1967); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Neotemnopteryx fulva Saussure (= Gislenia australica Brunner), Tp, Gb. NEW SOUTH WALES: 
Glen Dhu Cave (Allston Cave) (TR15), Timor (Richards 1967a); Murder Cave (CL2), Cliefden 
(Richards 1967a); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Richards 1967a); Haystall 
Cave (U23), Naracoorte (Richards 1967a); VICTORIA: Mabel Cave (EB1), East Buchan (Richards 
1967a). 

Neotemnopteryx sp., Tp, Gp?. NEW SOUTH WALES: Ashford Main Cave (AS.l), Ashford 
(Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe 
(Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith 
unpublished data); Johannsen's Cave (Jl-2), Limestone Ridge, Rockhampton (Hamilton-Smith 
unpublished data); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); 
Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Viator Main Cave (VR1), 
Viator Hill (Hamilton-Smith unpublished data). 

Weotemnopteryx (IGislenia sp.), Tp, Gp?. NEW SOUTH WALES: Ashford Main Cave (AS1), 
Ashford (Richards 1967a); Cave 4, Comboyne (Richards 1967a); Hill Cave (TR7), Timor (Richards 
1967a); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Richards 1967a); Swallow Cave 
(CU1), Cudgegong (Richards 1967a); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe 
(Richards 1967a); Riverton Main Cave (RN1), Riverton, southern Queensland (Richards 1967a); 
Viator Cave (VR4), Viator Hill, southern Queensland (Richards 1967a); Johannsen's Cave (Jl), 
Limestone Ridge, Rockhampton (Hamilton-Smith 1967); Winding Stairway Cave (4E2), Mt Etna 
(Hamilton-Smith 1967); SOUTH AUSTRALIA: Alexandra Cave (5U3), Naracoorte (Richards 
1967a); Bat Cave (U2), Naracoorte (Richards 1967a). 

Paratemnopteryx atra Princis, Tb, Gp. WESTERN AUSTRALIA: Mines near Marble Bar (Princis 
1963; Richards 1967a; Moore et al. 2001). 

Paratemnopteryx rufa Tepper, Gb?. NULLARBOR PLAIN: Murrawijinie No.3 Cave (N9) 
(Richards 1971); Abrakurrie Cave (N3) (Richards 1971). 

Paratemnopteryx sp., Tp, Gb?. QUEENSLAND: Pinwill Cave (4U17), Undara (Howarth 1988). 

Shawella douglasi Princis, Tp, Gb?. NEW SOUTH WALES: River Cave (SCI), Stockyard Creek 
(Hamilton-Smith 1967); WESTERN AUSTRALIA: Drovers Cave (J2), Jurien Bay (Hamilton-Smith 
unpublished data); Jurien Bay caves (Princis 1963; Richards 1967a); Eneabba Caves (El-3), 
Eneabba (Lowry 1996); Weelawadji Cave (E24), Eneabba (Lowry 1996). 

Trogloblattella nullarborensis Mackerras, Tb, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) 
(Mackerras 1967; Richards 1971); Koonalda Cave (N4) (Mackerras 1967); Mullamullang Cave 
(N37) (Mackerras 1967); Roaches Rest Cave (N58) (Mackerras 1967); Arubiddy Cave (N81) 
(Mackerras 1967). 

Blattidae 

Polyzosteria mitchelli Angas, Tp. NULLARBOR PLAIN: Warbla Cave (Nl), (Mackerras 1965); 
Kestrel Cavern (N40) (Mackerras 1965; Richards 1967a). 

Polyzosteria pubescens Tepper, Tp. NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton- 
Smith unpublished data). 

20 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 

Zonioploca medilinea Tepper, Tp?. NULLARBOR PLAIN: Warbla Cave (Nl) (Richards 1967a). 

Order Orthoptera 

Rhaphidophoridae 

Australotettix carraiensis Richards, Tp, Gx. NEW SOUTH WALES: Barnett's Cave (SC6), 
Stockyard Creek (Richards 1964); Carrai Bat Cave (SC5), Stockyard Creek (Richards 1964); Col's 
Cave, Stockyard Creek (Richards 1964); Lot's Mansion, Stockyard Creek (Richards 1964); River 
Cave (SCI) Stockyard Creek (Richards 1964). 

Cavernotettix buchanensis Richards, Tx, Gx. VICTORIA: Wilson Cave (EB4), East Buchan 
(Richards 1966; Yen and Milledge 1990); Trogdip Cave (EB10), East Buchan (Hamilton-Smith 
unpublished data); Spring Creek Cave (Bl), Buchan (Richards 1966; Yen and Milledge 1990); 
Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal 
(Yen and Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990); Dickson 
Cave (M30), Murrindal (Yen and Milledge 1990); Nargun's Cave (NN1), Nowa Nowa Caves 
(Richards 1966; Yen and Milledge 1990); Weta Cave (NN2), Nowa Nowa Caves (Richards 1966; 
Yen and Milledge 1990). 

Cavernotettix montanus Richards, Tx, Gx. NEW SOUTH WALES: small cave nr Glory Cave, 
Yarrangobilly (Richards 1966); Jersey Cave (Y23), Yarrangobilly (Richards 1966); Restoration 
Cave (Y50), Yarrangobilly (Richards 1966); Unnamed cave, Yarrangobilly (Richards 1966); 
Cooleman Cave (CP1), Cooleman Plains (Richards 1966); Unnamed cave opp. Blue Waterhole, 
Cooleman Plains (Richards 1966); Unnamed cave nr Murray Cave, Cooleman Plains (Richards 
1966). 

Cavernotettix wyanbenensis Richards, Tx, Gx. NEW SOUTH WALES: Wyanbene Cave (WY1), 
Wyanbene (Richards 1966); Bat Cave, Cheitmore (Richards 1966). 

Pallidotettix nullarborensis Richards, Tx, Gx. NULLARBOR PLAIN: Warbla Cave (Nl) (Richards 
1971); Weebubbie Cave (N2) (Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); 
Cocklebiddy Cave (N48) (Richards 1971); Pannikin Plain Cave (N49) (Richards 1971); Tommy 
Grahams Cave (N56) (Richards 1971). 

Undetermined genus and species, Tx, Gx. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); Colong Main Cave (CGI), Colong (Hamilton-Smith 
unpublished data); QUEENSLAND: Danes Four Cave (C4), Camooweal (Hamilton-Smith 
unpublished data); Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal 
(Hamilton-Smith unpublished data); Haunted Cave (CHI), Chillagoe (Hamilton-Smith unpublished 
data); VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds unpublished data). 

Order Psocoptera 

Liposcelidae 

Liposcelis corrodens Broadhead, Tpl, Gp. WESTERN AUSTRALIA: Arranmall [sic] Cave (E22), 
Eneabba (Smithers 1975); undetermined caves (Smithers 1975). 

Psyllipsocidae 

Wsyllipsocus ramburi Selys-Longcamp, Tpl, Gp. NEW SOUTH WALES: Murder Cave (CL2), 
Cliefden (Hamilton-Smith 1967); Island Cave (CL6), Cliefden (Smithers 1964); Hill Cave (TR7), 
Timor (James et al. 1976); Basin Cave (W4), Wombeyan (Smithers 1964); Fig Tree Cave (W148), 
Wombeyan (Smithers 1975); Punchbowl Cave (WJ8), Wee Jasper (Smithers 1964); Church Cave 
(WJ31), Wee Jasper (Smithers 1964); NULLARBOR PLAIN: Weebubbie Cave (N2) (Richards 
1971); Abrakurrie Cave (N3) (Hamilton-Smith 1967; Richards 1971); Koonalda Cave (N4) 
(Richards 1971); Madura Cave (N62), (Richards 1971); QUEENSLAND: Riverton Main Cave 
(RN1), Riverton, southern Queensland (Hamilton-Smith 1967); SOUTH AUSTRALIA: Bat Cave 



Proc. Linn. Soc. N.S.W., 125, 2004 21 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

(U2), Naracoorte (Smithers 1964; Bellati et al. 2003); Blackberry Cave, Naracoorte (Smithers 
1964); VICTORIA: Clogg's Cave (EB2), East Buchan (Smithers 1964); O'Rourkes Cave (B12), 
Buchan (Smithers 1964); Nargun's Cave (NN1), Nowa Nowa (Hamilton-Smith 1967). 

Trogiidae 

Lepinotus inquilinus Heyden, Tpl, Gp. WESTERN AUSTRALIA: Arranmall (sic) Cave (E22), 
Eneabba (Smithers 1975). 

ILepinotus reticulatus Enderlein, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Smithers 
1964; Bellati et al. 2003) 

Undetermined genus and species, NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan 
(Dennis and Mayhew 1986). 

Undetermined Family 

Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Gable Cave (CL7), Cliefden 
(Hamilton-Smith unpublished data); QUEENSLAND: Viator Main Cave (VR1), Viator Hill 
(Hamilton-Smith unpublished data); VICTORIA: Lilly Pilly Cave (M8), Murrindal (Yen and 
Milledge 1990). 

Order Hemiptera 

Cixiidae 

Undetermined genus and species, Tp. QUEENSLAND: Mount Etna Main Cave (El), Mount Etna 
(Hamilton-Smith unpublished data); Johannsen's Cave (Jl-2), Limestone Ridge, Rockhampton 
(Hamilton-Smith unpublished data). 

Lygaeoidea 

Undetermined family and genus, Gp?. VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds 
unpublished data). 

Reduviidae 

Armstrongula sp. Tp, Gp. SOUTH AUSTRALIA: McKinley's Daughter's Cave (F175), Flinders 
Ranges (T. Moulds unpublished data); Unnamed mine, Weetootla Gorge, Gammon Ranges (T. 
Moulds unpublished data). 

Centrogonus sp. Tp, Gp. NORTHERN TERRITORY: Kintore Cave (K2), Katherine (Hamilton- 
Smith unpublished data). 

Undetermined Emesinae genus and species, Tp, Gp. QUEENSLAND: Crazy Cracks Cave, Jacks 
Gorge, Broken River (T. Moulds unpublished data); Not Another Frig Tree Crave, Jacks Gorge, 
Broken River (T. Moulds unpublished data); Johannsen's Cave (Jl-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton 
(Hamilton-Smith unpublished data). 

Undetermined genus and species, Tp, Gp. NORTHERN TERRITORY: Cutta Cutta Cave (Kl), 
Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Queenslander Cave (CH15), 
Queenslander Tower (CH5246) Chillagoe (T. Moulds unpublished data); Trezkinn Cave (CH14), 
Chillagoe (T. Moulds unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith 
unpublished data). 

Undetermined genus and species, Tp, Gp?. QUEENSLAND: Elephant Hole (E8), Mount Etna 
(Hamilton-Smith unpublished data). 



22 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Undetermined Family 

Undetermined genus and species, QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton- 
Smith unpublished data). 

Order Neuroptera 

Myrmeleontidae 

Aeropteryx sp., Tp, Gp. SOUTH AUSTRALIA: McKinley's Daughter's Cave (F175), Flinders 
Ranges (T. Moulds unpublished data); Moro Bat Cave (F47), Flinders Ranges (T. Moulds 
unpublished data); Unnamed cave, Brachina Gorge, Flinders Ranges (T. Moulds unpublished data); 
Unnamed bat cave, Chambers Gorge, Flinders Ranges (T. Moulds unpublished data); Unnamed 
cave, Chambers Gorge, Flinders Ranges (T. Moulds unpublished data); Unnamed mine, Weetootla 
Gorge, Gammon Ranges (T. Moulds unpublished data). 

Myrmeleontinae sp., Tp?. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith 
unpublished data). 

Undetermined Family 

Undetermined genus and species, QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer's 
Mountain (Hamilton-Smith unpublished data). 

Order Coleoptera 

Anobiidae (Ptinidae) 

Ptinus exulans Erichson, Tpl, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith 1967); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Island 
Cave (CL6), Cliefden (Hamilton-Smith 1967); Jenolan Caves (Hamilton-Smith 1967); Willi Willi 
Bat Cave (WW1), Willi Willi (Hamilton-Smith 1967); Bungonia various caves (Eberhard 1998); 
Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Main Cave (CGI), Colong 
(Hamilton-Smith unpublished data); NULLARBOR PLAIN: Warbla Cave (Nl) (Richards 1971); 
Murrawijinie No. 1 Cave (N7) (Richards 1971); Murra-El-Elevyn Cave (N47) (Hamilton-Smith 
1967; Richards 1971); Firestick Cave (N70) (Richards 1971); Dingo Cave (N160) (Richards 1971); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 1967; Bellati et al. 2003); 
Blanche Cave (U4), Naracoorte (Hamilton-Smith 1967); VICTORIA: Starlight Cave (W5), 
Warrnambool (Hamilton-Smith 1967); Clogg's Cave (EB2), East Buchan (Hamilton-Smith 1967); 
WESTERN AUSTRALIA: Goosebury Cave (Jl), Jurien Bay (Hamilton-Smith 1967). 

Carabidae 

Anomotarus subterraneus Moore, Tp, Gp. QUEENSLAND: Riverton Main Cave (RN1), Riverton, 
southern Queensland (Moore 1967). 

Cratogaster melus Laporte, Tp?. QUEENSLAND: Johannsen's Cave (Jl-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data). 

Darodilia sp., Tp?. QUEENSLAND: Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith 
unpublished data). 

Gnathaphanus pulcher Dejean, Tp?. NORTHERN TERRITORY: Cutta Cutta Cave (Kl), Katherine 
(Hamilton-Smith unpublished data); Kintore Cave (K2), Katherine (Hamilton-Smith unpublished 
data); QUEENSLAND: Johannsen's Cave (Jl-2), Limestone Ridge, Rockhampton (Hamilton-Smith 
unpublished data). 

Lecanomerus sp., Gp?. NEW SOUTH WALES: Youndales Cave (Hut Cave) (KB1), Kunderang 
Brook (Hamilton-Smith unpublished data). 

Mecyclothorax ambiguus Erichson, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- 
Smith unpublished data). 

Proc. Linn. Soc. N.S.W., 125, 2004 23 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

Meonis sp., Tp, Gp. QUEENSLAND: Main Mount Etna Cave (El), Mount Etna (Hamilton-Smith 
unpublished data). 

Mystropomus subcostatus Chaudoir, Tp?. QUEENSLAND: Johannsen's Cave (J 1-2), Limestone 
Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna 
(Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith 
unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Piglet 
Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data). 

Notonomus angustibasis Sloane, Tp?, Gx. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 

Notospeophonus castaneus castaneus Moore, Tp2. SOUTH AUSTRALIA: Blanche Cave (U4), 
Naracoorte (Hamilton-Smith 1967); Blackberry Cave (U8), Naracoorte (Hamilton-Smith 1967); 
Stick Cave (Ull), Naracoorte (Moore 1964); Cathedral Cave (U12), Naracoorte (Moore 1964); Fox 
Cave (U22), Naracoorte (Hamilton-Smith 1967); Haystall Cave (U23), Naracoorte (Hamilton-Smith 
1967); Cave Park Cave (U37), Naracoorte (Hamilton-Smith unpublished data); Tantanoola Caves 
(Hamilton-Smith 1967); VICTORIA: Bat Cave (P6), Portland (Moore 1962); Byaduk Caves, 
Byaduk (Moore 1962); Panmure Cave (H5), Mount Napier (Moore 1964); Mt Widderin Cave (HI), 
Skipton (Hamilton-Smith 1967); Snowflake Cave (LI), Glenelg River (Hamilton-Smith 1967); 
Curran's Creek Cave (G4), Glenelg River (Hamilton-Smith 1967). 

Notospeophonus castaneus consobrinus Moore, Tp, Gp. VICTORIA: Spring Creek Cave (Bl), 
Buchan (Hamilton-Smith unpublished data); Moon Cave (B2), Buchan (Hamilton-Smith 
unpublished data); Mabel Cave (EB1), East Buchan (Hamilton-Smith unpublished data); Wilson's 
Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip Cave (EB10), East Buchan 
(Hamilton-Smith unpublished data); Slocombe's Cave (BA1), The Basin (Hamilton-Smith 
unpublished data); Shades of Death Cave (M3), Murrindal (Hamilton-Smith unpublished data); 
Anticline Cave (Mil), Murrindal (Hamilton-Smith unpublished data); SSS Cave (M44), Murrindal 
(Hamilton-Smith unpublished data). 

Notospeophonus jasper ensis jasper ensis Moore, Tp2, Gp. NEW SOUTH WALES: Punchbowl Cave 
(WJ8), Wee Jasper (Moore 1964); Pylon 58 Cave (WJ99), Wee Jasper (Moore 1964); Basin Cave 
(W4), Wombeyan (Hamilton-Smith unpublished data). 

Notospeophonus jasperensis vicinus Moore, Tp2, Gp. NEW SOUTH WALES: Bungonia various 
caves (Eberhard 1998). 

Notospeophonus pallidus Moore, Tp2, Gp?. NEW SOUTH WALES: Childrens Cave (CL12), 
Cliefden (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Myponga (Moore 1964); 
NULLARBOR PLAIN: Warbla Cave (Nl) (Hamilton-Smith 1967; Richards 1971); Weebubbie 
Cave (N2) (Richards 1971); Abrakurrie Cave (N3) (Hamilton-Smith 1967; Richards 1971); 
Koonalda Cave (N4) (Hamilton-Smith 1967; Richards 1971); Koomooloobooka Cave (N6) 
(Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); Knowles Cave (N22) (Hamilton- 
Smith 1967; Richards 1971); Mullamullang Cave (N37) (Richards 1971); Joe's Cave (N39) 
(Hamilton-Smith 1967; Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Madura Cave 
(Madura 6 Mile South Cave) (N62) (Richards 1971); Lynch Cave (N60) (Richards 1971). 

Notospeophonus sp., Tp, Gp?. QUEENSLAND: Viator Main Cave (VR1), Viator Hill (Hamilton- 
Smith unpublished data). 

Phloeocarabus sp. Tp?, Gp?. QUEENSLAND: Haunted Cave (CHI), Chillagoe (Hamilton-Smith 
unpublished data). 



24 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 

Pogonoglossus sp., Tp, Gp?. NORTHERN TERRITORY: Cutta Cutta Cave (Kl), Katherine 
(Hamilton-Smith unpublished data). 

Pseudoceneus sp. Tp, Gp?. WESTERN AUSTRALIA: Stockyard Cave (E3), Eneabba (Hamilton- 
Smith unpublished data). 

Speotarus lucifugus Moore, Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Moore 
1964; Bellati et al. 2003); NULLARBOR PLAIN: Warbla Cave (Nl) (Richards 1971); Weebubbie 
Cave (N2) (Richards 1971); Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 
1971); Winbirra Cave (N45) (Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); 
Cocklebiddy Cave (N48) (Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Lynch 
Cave (N60) (Richards 1971); Unnamed cave (N139) (Richards 1971). 

Speotarus princeps Moore, Tp2, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Moore 1964); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished 
data). 

Speotarus sp., Tp, Gp. NULLARBOR PLAIN: Warbla Cave (Nl) (Hamilton-Smith unpublished 
data); Weebubbie Cave (N2) (Hamilton-Smith unpublished data); Murra-El-Elevyn Cave (N47) 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Mount Sims Cave (F7), Walpunda 
Creek, Flinders Ranges (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Gooseberry 
Cave (Jl), Jurien Bay (Hamilton-Smith unpublished data). 

Thenarotes speluncarius Moore, Tp, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 
1971); Koonalda Cave (N4) (Richards 1971); New Cave (Nil) (Richards 1971); Lynch Cave (N60) 
(Richards 1971); Decoration Cave (N84) (Richards 1971); SOUTH AUSTRALIA: Cave No. 1, 
Buckalowie, Flinders Ranges (Hamilton-Smith unpublished data). 

Trechimorphus diemenensis Bates, Tpl, Gx. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Jenolan Caves 
(Moore 1964); VICTORIA: Dalley's Sinkhole (M35), Murrindal (Hamilton-Smith 1967). 

Trichosternus vigorsi Gory, Tp? Gx. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 

Undetermined genus and species, NEW SOUTH WALES: Grill Cave (B44), Bungonia (Eberhard 
and Spate 1995); Belfry Cave (TR2), Timor (James et al. 1976); Glen Dhu Cave (Allston Cave) 
(TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (Tl), Tuglow (Hamilton-Smith 
unpublished data); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), 
Camooweal (Hamilton-Smith unpublished data); Mount Etna Main Cave (El), Mount Etna 
(Hamilton-Smith unpublished data); Cave with the thing that went thump! (E5), Mount Etna 
(Hamilton-Smith unpublished data). 

Undetermined genus and species, Tp, Gp. QUEENSLAND: Barker's Cave (U34), Undara 
(Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (Bl), Buchan (Yen and 
Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson's Cave (EB4), 
East Buchan (Hamilton-Smith unpublished data); Shades of Death Cave (M3), Murrindal (Yen and 
Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990). 

Cryptophagidae 

Anchicera sp., Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Atomaria sp., Gp. Southern Australia (Hamilton-Smith 1968). 



Proc. Linn. Soc. N.S.W., 125, 2004 25 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Basin Cave (W4), Wombeyan 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Fox Cave (U22), Naracoorte 
(Hamilton-Smith unpublished data); VICTORIA: Wilson's Cave (EB4), East Buchan (Hamilton- 
Smith unpublished data); Nargun's Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data). 

Curculionidae 

Mandalotus sp. Gp?. NEW SOUTH WALES: Chalk Cave (B26), Bungonia (Hamilton-Smith 
unpublished data). 

Talaurinus sp. Gp?. QUEENSLAND: Johannsen's Cave (J 1-2), Mount Etna (Hamilton-Smith 
unpublished data). 

Dermestidae 

Dermestes ater DeGeer, Tp, Gp. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton- 
Smith unpublished data). 

Undetermined genus and species, Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Bellati et al. 2003); QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer's Mountain (Hamilton- 
Smith unpublished data); Unidentified cave in southern Queensland (Hamilton-Smith 1967); 
VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). 

Endomychidae 

Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith unpublished data). 

Histeridae 

Carcinops sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) 
(Humphreys and Eberhard 2001). 

Saprinus sp., Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith 
unpublished data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith 
unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 
unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished 
data); Nargun's Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Clogg's Cave 
(EB2), East Buchan (Hamilton-Smith unpublished data). 

Tomogenius Iripicola Marseul, Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati 
et al. 2003); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); 
NULLARBOR PLAIN: Lynch Cave (N60) (Richards 1971); Thylacine Hole (N63) (Richards 
1971); Dingo Cave (N160) (Richards 1971). 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998); Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); Carrai 
Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main 
Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); QUEENSLAND: Holy Jump Lava 
Cave (BM1), Bauer's Mountain (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), 
Riverton (Hamilton-Smith unpublished data); Johannsen's Cave (Jl-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Sand Cave (Joanna) (U16), Naracoorte 
(Hamilton-Smith unpublished data); VICTORIA: Chimney Cave (BR1), Bat Ridges, Portland 
(Hamilton-Smith unpublished data); Clogg's Cave (EB2), East Buchan (Hamilton-Smith 
unpublished data); Nargun's Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Bat 
Cave (P6), Portland (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Gooseberry 
Cave (Jl), Jurien Bay (Hamilton-Smith unpublished data). 



26 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Jacobsoniidae 

Derolathrus sp., Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); 
Various caves in southern Australia (Hamilton-Smith 1967). 

Undetermined genus and species, Tp, Gb. VICTORIA: Bat Cave (P6), Portland (Hamilton-Smith 
unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 

Lathridiidae 

Corticaria sp., Gp. Southern Australia (Hamilton-Smith 1968); NEW SOUTH WALES: Ashford 
Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie 
Cave (N2) (Hamilton-Smith unpublished data); Abrakurrie Cave (N3) (Hamilton-Smith unpublished 
data): VICTORIA: Skipton Cave (Mount Widderin Cave) (HI), Mount Napier (Hamilton-Smith 
unpublished data). 

Leiodidae 

Choleva austmlis, Tp, Gp. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith 
unpublished data). 

Choleva sp., Tp, Gp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Richards 1971); Lynch 
Cave (N60) (Richards 1971). 

Nargomorphus minusculus Blackburn, Tpl, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Hamilton-Smith 1967; Bellati et al. 2003); VICTORIA: Anticline Cave (Mil), Murrindal 
(Hamilton-Smith 1967). 

Pseudonemadus adelaidae Blackburn, Tp, Gp. NEW SOUTH WALES: Glen Dhu Cave (Allston 
Cave) (TR15), Timor (Hamilton-Smith unpublished data); QUEENSLAND: Riverton Main Cave 
(RN1), Riverton (Hamilton-Smith unpublished data). 

Pseudonemadus australis Erichson, Gp. VICTORIA: Chimney Cave (BR1), Bat Ridge, Portland 
(Hamilton-Smith unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); 
Panmure Cave (H5), Mt Napier (Hamilton-Smith unpublished data). 

Pseudonemadus integer Portevin, Gp. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data); QUEENSLAND: Speaking Tube (E7), Mount Etna (Hamilton- 
Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Mt Widderin 
Cave (HI), Skipton (Hamilton-Smith unpublished data); Panmure Cave (H5), Mt Napier (Hamilton- 
Smith unpublished data). 

Pseudonemadus sp., Gp. Southern Australia (Hamilton-Smith 1968). 

?Leiodidae 

Undetermined genus and species, NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Smith 
1982a). 

Melyridae 

Heteromastix sp. Tx?, Gx?. NEW SOUTH WALES: Colong Main Cave (CGI), Colong (Hamilton- 
Smith unpublished data). 

Merophysiidae 

Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith 1967). 



Proc. Linn. Soc. N.S.W., 125, 2004 27 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Pselaphidae 

Rybaxisl sp., Tp, Gp. NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Hamilton-Smith 
1966a); Bungonia various caves (Eberhard 1998). 

Tyromorphus speciosus King, Tpl. NEW SOUTH WALES: Unidentified cave, Southern 
Limestone, Jenolan (Hamilton-Smith 1966a); Paradox Cave (J48), Jenolan (Hamilton-Smith 
unpublished data); QUEENSLAND: Johannsen's Cave (Jl-2), Limestone Ridge, Rockhampton 
(Hamilton-Smith 1966a); VICTORIA: Anticline Cave (Mil), Murrindal (Hamilton-Smith 1966a). 

Undetermined genus and species, Gp. QUEENSLAND: Rope Ladder Cave, Mingella (Weinstein 
and Slaney 1995). 

Undetermined genus and species, Tp, Gp. VICTORIA: Wilson's Cave (EB4), East Buchan 
(Hamilton-Smith unpublished data). 

Ptilidae 

Achosia lanigera Deane, Tp?, Gp. VICTORIA: Wilsons Cave (EB4), East Buchan (Hamilton-Smith 
unpublished data). 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 

Rhizophagidae 

Undetermined genus and species, Gp. QUEENSLAND: Rope Ladder Cave (FR2), Mingella, 
Fanning River (Weinstein and Slaney 1995). 

Scarabaeidae 

Aulacopris maximus Matthews, Tpl, Gb. NEW SOUTH WALES: Yessabah Bat Cave (YE1), 
Yessabah (Waite 1898); Unknown cave in Coorabakh National Park (formerly part Lansdowne 
State Forest), Taree (Williams 2003). 

Aulacopris reichei White, Tpl, Gp. NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah 
(Lea 1923); Unknown cave, Mosman (Fricke 1964). 

Amphistomus accidatus Matthews, Tx, Gp. QUEENSLAND: Elephant Hole (E8), Mount Etna 
(Hamilton-Smith unpublished data). 

Saprosites mendax Blackburn, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte 
(Hamilton-Smith unpublished data). 

Undetermined genus and species, Gp. NEW SOUTH WALES: Willi Willi Bat Cave (Main Cave) 
(WW1), Willi Willi (Hamilton-Smith unpublished data). 

Silphidae 

Ptomaphila lachrymosa Schreibers, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- 
Smith unpublished data). 

Staphylinidae 

Myotyphlus jansoni Matthews, Tpl, Gp. NEW SOUTH WALES: Unidentified cave, Southern 
Limestone, Jenolan (Hamilton-Smith and Adams 1966); Paradox Cave (J48), Jenolan (Hamilton- 
Smith unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith and 
Adams 1966); Bat Cave (P6), Portland (Hamilton-Smith 1967). 



28 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 

Philonthus parous Sharp, Gp?. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(Moore 1964); Church Cave (WJ31), Wee Jasper (Moore 1964); VICTORIA: Starlight Cave (W5), 
Warrnambool (Hamilton-Smith unpublished data). 

Quedius luridipennis Macleay, Tp?, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 
1971). 

Quedis sp., Gp. Southern Australia (Hamilton-Smith 1968); NEW SOUTH WALES: Church Cave 
(WJ31), Wee Jasper (Hamilton-Smith unpublished data); Signature Cave (WJ7) (Hamilton-Smith 
unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 
unpublished data); VICTORIA: Wilsons Cave (EB4), East Buchan (Hamilton-Smith unpublished 
data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998); Colong Main Cave (CGI), Colong (Hamilton-Smith unpublished data); Dip Cave 
(WJ1), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper 
(Hamilton-Smith unpublished data); NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton- 
Smith unpublished data); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum 
Cave), Camooweal (Hamilton-Smith unpublished data); Camooweal Four Mile East Cave 
(Camooweal Cave, Four Mile Cave) (CI 3), Camooweal (Hamilton-Smith unpublished data); 
VICTORIA: Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Grassmere Cave (W6), 
Warrnambool (Hamilton-Smith unpublished data). 

Undetermined genus and species , Tp?, Gx?. QUEENSLAND: Four Mile Cave (C14), Camooweal 
(Hamilton-Smith unpublished data). 

Tenebrionidae 

Adelium sp. Tp? Gp?. SOUTH AUSTRALIA: Sand Cave (Joanna) (U16), Naracoorte (Hamilton- 
Smith unpublished data). 

Alphitobius laevigatus Fabricius, Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux 
Cave (CI56) (Humphreys and Eberhard 2001). 

Alphitobius diaperinus Panzer, Tp?, Gp?. QUEENSLAND: Bat Cleft (E6), Mount Etna (Hamilton- 
Smith unpublished data). 

Brises acuticornis Pascoe, Tpl, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith 1967); NORTHERN TERRITORY: Unknown bat caves, 15 km south of Alice 
Springs (Mathews 1986); NULLARBOR PLAIN: Warbla Cave (Nl) (Richards 1971); Weebubbie 
Cave (N2) (Hamilton-Smith 1967; Richards 1971); Abrakurrie Cave (N3) (Richards 1971); 
Koonalda Cave (N4) (Richards 1971); Koomooloobooka Cave (N6) (Richards 1971); Murrawijinie 
No. 1 Cave (N7) (Richards 1971); Murrawijinie No. 2 Cave (N8) (Richards 1971); Murrawijinie 
No.3 Cave (N9) (Richards 1971); White Wells Cave (N14) (Hamilton-Smith 1967; Richards 1971); 
Unnamed cave (N33) (Mathews 1986); Mullamullang Cave (N37) (Hamilton-Smith 1967; Richards 
1971); Winbirra Cave (N45) (Richards 1971); Nurina Cave (N46) (Richards 1971); Murra-El- 
Elevyn Cave (N47) (Hamilton-Smith 1967; Richards 1971); Cocklebiddy Cave (N48) (Richards 
1971); Pannikin Plain Cave (N49) (Richards 1971); Moonera Tank Cave (N53) (Richards 1971); 
Tommy Grahams Cave (N56) (Richards 1971); Lynch Cave (N60) (Richards 1971); White Wells 
Blowhole (N61) (Hamilton-Smith 1967); Madura Cave (Madura Six Miles South Cave) (N62) 
(Hamilton-Smith 1967; Richards 1971); Thylacine Hole (N63) (Richards 1971); Firestick Cave 
(N70) (Richards 1971); Old Homestead Cave (N83) (Richards 1971); Diprose No.l Cave (N96) 
(Hamilton-Smith 1967); Diprose No.3 Cave (N98) (Hamilton-Smith 1967); Snake Pit (N133) 
(Richards 1971); Unnamed cave (N149) (Richards 1971); Dingo Cave (N160) (Richards 1971); 
Swallow Cave, Cocklebiddy (Mathews 1986); SOUTH AUSTRALIA: Punyelroo Cave (Ml), Swan 



Proc. Linn. Soc. N.S.W., 125, 2004 29 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

Reach Murray Plains (Hamilton-Smith 1967); Clara St. Dora Cave (F4), Buckalowie, Flinders 
Ranges (Hamilton-Smith 1967); Unknown bat cave, Wilpena Pound, Flinders Ranges (Mathews 
1986); Wooltana Cave (F9), Flinders Ranges (Mathews 1986). 

Brises katherinae Matthews, Tp, Gp. NORTHERN TERRITORY: Cutta Cutta Cave (Kl), Katherine 
(Mathews 1986; Hamilton-Smith et al. 1989); Kintore Cave (K2), Katherine (Mathews 1986); Three 
Mile Cave, Katherine (Mathews 1986); 16 Mile Cave, Katherine (Mathews 1986). 

Helea catenulatusl Mail., Gp?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton- 
Smith unpublished data). 

Pterohelaeus piceus Kirby, Tp, Gp?. NEW SOUTH WALES: Island Cave (CL6), Cliefden 
(Hamilton-Smith unpublished data); Timor Caves, Timor (Hamilton-Smith unpublished data). 

Pterohelaeus sp., Tp, Gp?. NEW SOUTH WALES: Main Cave (Ballroom Cave) (TR1), Timor 
(Hamilton-Smith unpublished data); QUEENSLAND: Johannsen's Cave (Jl-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data); Ilium Cave (E31), Mount Etna (Hamilton-Smith 
unpublished data). 

Undetermined genus and species, Gp. NEW SOUTH WALES: Colong Main Cave (CGI), Colong 
(Hamilton-Smith unpublished data); Moparabah Cave (MP1), Moparabah (Hamilton-Smith 
unpublished data). 

Trogidae 

Omorgus costatus Wiedemann, Tp, Gp?. QUEENSLAND: Johannsen's Cave (Jl-2), Limestone 
Ridge, Rockhampton (Hamilton-Smith unpublished data). 

Trox alatus Macleay, Tp, Gp. NORTHERN TERRITORY: various caves of the Kimberley region 
(Hamilton-Smith et al. 1989); Kintore Cave (K2), Katherine (Hamilton-Smith unpublished data). 

Trox amictus Haaf, Tp, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971); 
Murrawijinie No. 1 Cave (N7) (Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); 
Mullamullang Cave (N37) (Richards 1971); Lynch Cave (N60) (Richards 1971); Skink Hole (N82) 
(Richards 1971); Old Homestead Cave (N83) (Richards 1971); Decoration Cave (N84) (Richards 
1971). 

Trox sp., Tp, Gp. NEW SOUTH WALES: guano caves from northern NSW (Hamilton-Smith 
1967); Comboyne C4 Cave, Comboyne (Hamilton-Smith unpublished data); NORTHERN 
TERRITORY: various bat caves (Hamilton-Smith 1967); QUEENSLAND: Riverton Main Cave 
(RN1), Riverton (Hamilton-Smith unpublished data); Arch Cave (U22), Undara (Hamilton-Smith 
unpublished data); Barker's Cave (U34), Undara (Hamilton-Smith unpublished data); various bat 
caves (Hamilton-Smith 1967); Bat Cleft (E6), Mount Etna (Hamilton-Smith unpublished data). 

Undetermined Family 

Undetermined genus and species, Tp, Gp. QUEENSLAND: Clam Cave (CH26), Walkunder Tower, 
Chillagoe (Matts 1987); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); 
Bat Cleft (E6), Mount Etna (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave 
(Bl), Buchan (Yen and Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 
1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). 

Order Siphonaptera 

Ischnopsyllidae 

Porribius sp., Tx, Gx, P. NULLARBOR PLAIN: Mullamullang Cave (N37) (Richards 1971); 
Warbla Cave (Nl) (T. Moulds unpublished data). 



30 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 

Order Diptera 

Anthomyiidae 

Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith unpublished data); Southern Australia (Hamilton-Smith 1968). 

Cecidomyidae 

Undetermined genus and species, Gp. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith 
unpublished data). 

Ceratopogonidae 

Culicoides sp., Tp, Gx?. QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum 
Cave), Camooweal (Hamilton-Smith unpublished data). 

Cypselosomatidae 

Clisa (Cypselosoma) australe McAlpine, Tpl, Gb. NEW SOUTH WALES: Carrai Bat Cave (SC5), 
Stockyard Creek (McAlpine 1966, 1993). 

Chloropidae 

Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux 
Cave (CI56) (Humphreys and Eberhard 2001). 

Chironomidae 

Diplocladius multiserialis Freeman, Tx, Gx?. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Hamilton-Smith unpublished data). 

Podonomus sp., Tx, Gx?. NEW SOUTH WALES: Mammoth Cave (J13), Jenolan (Hamilton-Smith 
unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data). 

Polypedilum watsoni Freeman, Tx, Gx?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Hamilton-Smith unpublished data). 

Tanytarus sp. Tx, Gx?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 
unpublished data). 

Undetermined genus and species,Tx, Gx?. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith 
unpublished data); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); 
QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal 
(Hamilton-Smith unpublished data). 

Chyromyidae 

Aphaniosoma sp., Tp, Gp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(McAlpine 1966); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); 
QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data). 

Dolicopodidae 

Sympycnus sp., Tx?, Gx?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith 
unpublished data). 

?Drosophilidae 

Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux 
Cave (CI56) (Humphreys and Eberhard 2001). 



Proc. Linn. Soc. N.S.W., 125, 2004 31 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Fanniidae 

Fannia sp., Tp, Gx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished 
data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 

Milichiidae 

Undetermined genus and species, Tp? Gx?. NEW SOUTH WALES: Church Cave (W31), Wee 
Jasper (Hamilton-Smith unpublished data); QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer's 
Mountain (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton- 
Smith unpublished data). 

Muscidae 

Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Grimes Cave (CI53) 
(Humphreys and Eberhard 2001); Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 
2001); NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished 
data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Bat Cave (P6), Portland (Hamilton-Smith unpublished data). 

Undetermined genus and species, Gx?. QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, 
Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data). 

Mycetophilidae 

Exechia pullicauda Skuse, Tp?, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data). 

Exechia sp., Tp?, Gp?. NEW SOUTH WALES: The Drum Cave (B13), Bungonia (Hamilton-Smith 
unpublished data). 

Nycteribiidae 

Basilia halei Musgrave, Tp. Gx, P. WESTERN AUSTRALIA: Wilgie-Mia Ochre Mine (MIS34), 
Cue (Hamilton-Smith unpublished data). 

Basilia musgravei Theodor, Tp, Gx, P. QUEENSLAND: Chillagoe Caves (Maa 1971); Pink's Cave 
(CH20), Chillagoe (Maa 1971); Viator Cave (VR4), Viator Hill (Maa 1971); Mount Etna Main 
Cave (El), Mount Etna (Maa 1971). 

Basilia techna Maa, Tp, Gx, P. NORTHERN TERRITORY: Red Bank Mine (Maa 1971); 
QUEENSLAND: Lawn Hill (Maa 1971); Olsen's Caves (Maa 1971). 

Basilia troughtoni Musgrave, Tp, Gx, P. WESTERN AUSTRALIA: Brown Bone Cave (SH17), 
South Hill River (Hamilton-Smith unpublished data). 

Basilia sp., Tp, Gx, P. QUEENSLAND: Tea-Tree Cave (CH43), Chillagoe (Hamilton-Smith 
unpublished data). 

Eremoctenia vandeuseni Maa, Tp, Gx, P. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Joanna Bat Cave (U38), Naracoorte 
(Hamilton-Smith unpublished data). 

Nycteribia allotopa meridiana Maa, Tp, Gx, P. QUEENSLAND: Pink's Cave (CH20), Chillagoe 
(Maa 1971); Mount Etna Main Cave (El), Mount Etna (Maa 1971). 

Nycteribia alternata Maa, Tp, Gx, P. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Maa 1971); Back Creek Mine (Maa 1971); Bonalbo Colliery (Maa 1971); Rise and Shine Mine 
(Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); QUEENSLAND: Phoenician 



32 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Mine (Maa 1971); Pink's Cave (CH20), Chillagoe (Maa 1971); Chillagoe Caves (Maa 1971); 
Mount Etna Main Cave (El), Mount Etna (Maa 1971). 

Nycteribia parilis vicaria Maa, Tp, Gx, P. NEW SOUTH WALES: Ashford Main Cave (AS1), 
Ashford (Maa 1971); Back Creek Mine (Maa 1971); Bonalbo Colliery (Maa 1971); Bullio Cave 
(W2), Wombeyan (Maa 1971); Carrai Bat Cave (SC5), Stockyard Creek (Maa 1971); Chietmore 
Cave (Maa 1971); Colong Caves (Maa 1971); Endless Cave (Maa 1971); Fig Tree Cave (W148), 
Wombeyan (Maa 1971); Gable Cave (CL7), Cliefden (Maa 1971); North Sydney railway tunnel 
(Maa 1971); Piano Cave (WA12), Walk' (Maa 1971); Prospect Tunnel (Maa 1971); Puchbowl Cave 
(WJ8), Wee Jasper (Maa 1971); Wee Jasper Caves (Maa 1971); Rise and Shine Mine (Maa 1971); 
Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); Wombeyan Caves (Maa 1971); Yessabah Bat 
Cave (YE1), Yessabah (Maa 1971); Bungonia various caves (Maa 1971; Hamilton-Smith 1972; 
Eberhard 1998); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); QUEENSLAND: 
Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith unpublished data); Pink's Cave (CH20), 
Chillagoe (Maa 1971); Tea-Tree Cave (CH43), Chillagoe (Hamilton-Smith unpublished data); 
Chillagoe Caves (Maa 1971); Phoenician Mine (Maa 1971); Pilkington Cave, Rockhampton (Maa 
1971); Mount Etna Main Cave (El), Mount Etna (Maa 1971); Cowie Bay Cave, Cape York 
Peninsula (Maa 1971); SOUTH AUSTRALIA: Hodges Cave (Joanna Bat Cave) (U38), Naracoorte 
(Maa 1971); Tomato-Stick Cave (U10), Naracoorte (Maa 1971); Bat Cave (U2), Naracoorte 
(Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (Bl), Buchan (Maa 1971); 
Grassmere Cave (W6), Warrnambool (Maa 1971); Mabel Cave (EB1), East Buchan (Hamilton- 
Smith unpublished data). 

Penicillidia oceanica Bigot, Tp, Gx, P. NEW SOUTH WALES: Ashford Main Cave (AS1), 
Ashford (Maa 1971); Back Creek Mine (Maa 1971); Belfery [sic] Cave (TR2), Timor (Maa 1971); 
Bonalbo Colliery (Maa 1971); Carrai Bat Cave (SC5), Stockyard Creek (Maa 1971); Chietmore 
Cave (Maa 1971); Colong Main Cave (CG3), Colong (Maa 1971); Drum Cave (B13), Bungonia 
(Maa 1971); Endless Cave (Maa 1971); Fig Tree Cave (W148), Wombeyan (Maa 1971); Gable 
Cave (CL7), Cliefden (Maa 1971); Prospect Tunnel (Maa 1971); Punchbowl Cave (WJ8), Wee 
Jasper (Maa 1971); Rise and Shine Mine (Maa 1971); Timor Main Cave (TR1), Timor (Maa 1971); 
Waterfall Gold Mine (Maa 1971); Yessabah Bat Cave (YE1), Yeassabah (Maa 1971); Bungonia 
various caves (Maa 1971; Hamilton-Smith 1972; Eberhard 1998); Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); QUEENSLAND: Chillagoe (Maa 1971); Holy Jump Lava 
Cave (BM1), Bauer's Mountain (Maa 1971); Johannsen's Cave (Jl-2), Limestone Ridge, 
Rockhampton (Maa 1971); VICTORIA: Greenhouse Cave (B3), Buchan (Maa 1971); Spring Creek 
Cave (Bl), Buchan (Hamilton-Smith unpublished data). 

Penicillidia setosala Maa, Tp, Gx, P. NEW SOUTH WALES: Fingal Point Cave (Maa 1971); Willi 
Willi Bat Cave (WW1), Willi Willi (Maa 1971); QUEENSLAND: Phoenician Mine (Maa 1971). 

Penicillidia tectisentis Maa, Tp, Gx, P. NEW SOUTH WALES: Willi Willi Bat Cave (WW1), Willi 
Willi (Maa 1971); QUEENSLAND: Mount Etna Main Cave (El), Mount Etna (Maa 1971); SOUTH 
AUSTRALIA: Tomato-Stick Cave (U10), Naracoorte (Maa 1971); Bat Cave (U2), Naracoorte 
(Hamilton-Smith unpublished data); VICTORIA: Grassmere Cave (W6), Warrnambool (Maa 1971). 

Penicillidia vandeuseni Maa, Tp, Gx, P. NEW SOUTH WALES: Fingal Point Cave (Maa 1971); 
Rise and Shine Mine (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); 
QUEENSLAND: Chillagoe Caves (Maa 1971); Royal Arch Cave (CH9), Chillagoe (Maa 1971); 
Tea-Tree Cave (CH43), Chillagoe (Hamilton-Smith unpublished data); Mount Etna Main Cave 
(El), Mount Etna (Maa 1971); SOUTH AUSTRALIA: Hodges Cave (Maa 1971). 

Phthiridium curvatum Theodor, Tp, Gx, P. NEW SOUTH WALES: Bonalbo Colliery (Maa 1971); 
Bullio Cave (W2), Wombeyan (Maa 1971); Cliefden (Maa 1971); Junction Cave (W152), 
Wombeyan (Maa 1971); Rise and Shine Mine (Maa 1971); Tanja Gold Mine (Maa 1971); 



Proc. Linn. Soc. N.S.W., 125, 2004 33 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

Morapabah Cave (Temagog Cave) (MP1), Morapabah (Maa 1971); Willi Willi Bat Cave (WW1), 
Willi Willi (Maa 1971); QUEENSLAND: Mount Etna Main Cave (El), Mount Etna (Maa 1971); 
VICTORIA: Mabel Cave (EB1), East Buchan (Maa 1971). 

Undetermined genus and species, Tp, Gx, P. QUEENSLAND: Flogged Horse Cave (Cammoo 
Cave) (J83), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); SOUTH 
AUSTRALIA: Asbestos mine near Arkaba, Flinders Ranges (Hamilton-Smith unpublished data); 
Drop Drop Cave (L29), Lower south east (Hamilton-Smith unpublished data). 

Phoridae 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998); The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); Colong 
Main Cave (CG3), Colong (Eberhard and Spate 1995); River Cave (CP6), Cooleman Plains 
(Eberhard and Spate 1995); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith 
unpublished data); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data); Urinary Tract 
Cave (W78), Wombeyan (Eberhard and Spate 1995); Fig Tree Cave (W148), Wombeyan 
(Hamilton-Smith unpublished data); Signature Cave (WJ7), Wee Jasper (Hamilton-Smith 
unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Dogleg 
Cave (WJ10), Wee Jasper (Eberhard 1993); Humicrib Cave (WJ34), Wee Jasper (Eberhard 1993); 
Pylon 58 Cave (WJ99), Wee Jasper (Eberhard 1993); NULLARBOR PLAIN: Abrakurrie Cave (N3) 
(Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); QUEENSLAND: Kaiser Creek 
Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); Cathedral Cave (U12), 
Naracoorte (Hamilton-Smith unpublished data); Fox Cave (U22), Naracoorte (Hamilton-Smith 
unpublished data); VICTORIA: Wilson's Cave (EB4), East Buchan (Hamilton-Smith unpublished 
data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). 

Platypezidae 

Undetermined genus and species, Gp. NEW SOUTH WALES: The Drum Cave (B13), Bungonia 
(Hamilton-Smith unpublished data). 

Psychodidae 

Phlebotumus sp., Tp, Gx?. QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum 
Cave), Camooweal (Hamilton-Smith unpublished data). 

Sergentomyia queenslandi Quate, Tp, Gx?. QUEENSLAND: Haunted Cave (CHI), Chillagoe 
(Hamilton-Smith unpublished data). 

Sergentomyia sp., Tp, Gx?. QUEENSLAND: Haunted Cave (CHI), Chillagoe (Hamilton-Smith 
unpublished data); Donna Cave (CH2), Chillagoe (Hamilton-Smith unpublished data); Royal Arch 
Cave (CH9), (Hamilton-Smith unpublished data); Royal Arch Cave (CH9), Chillagoe (Hamilton- 
Smith unpublished data); Trezkinn Cave (CH14), Chillagoe (Hamilton-Smith unpublished data); 
Keef s Cave (CH24), Chillagoe (Hamilton-Smith unpublished data); Tea-Tree Cave (CH43), 
Chillagoe (Hamilton-Smith unpublished data); Johannsen's Cave (J 1-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data). 

Undetermined genus and species, Tp. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et 
al. 2003); VICTORIA: Cloggs Cave (EB2), East Buchan (Hamilton-Smith unpublished data). 

Sciaridae 

Bradysia sp., Tp, Gp?. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton-Smith 
unpublished data); Willi Willi Bat Cave (WW1), Willi Willi (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); Cathedral 
Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: Panmure Cave (H5), 

34 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Mount Napier (Hamilton-Smith unpublished data). 

Lycoriella sp., Tp?, Gp?. NEW SOUTH WALES: Signature Cave (WJ7), Wee Jasper (Hamilton- 
Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Cloggs Cave (EB2), East Buchan (Hamilton-Smith unpublished data). 

Undetermined genus and species, Tp, Gp?. NEW SOUTH WALES: The Drum Cave (B13), 
Bungonia (Hamilton-Smith unpublished data); Chalk Cave (B26), Bungonia (Hamilton-Smith 
unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Main 
Cave (CGI), Colong (Hamilton-Smith unpublished data); Paradox Cave (J48), Jenolan (Hamilton- 
Smith unpublished data); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished 
data); Signature Cave (WJ7), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave 
(WJ8), Wee Jasper (Hamilton-Smith unpublished data); Church Cave (WJ31), Wee Jasper 
(Hamilton-Smith unpublished data); WiUi Willi Bat Cave (Main Cave) (WW1), Willi Willi 
(Hamilton-Smith unpublished data); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished 
data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith unpublished data); NULLARBOR 
PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith unpublished data); QUEENSLAND: Kaiser Creek 
Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003);VICTORIA: Clogg's Cave 
(EB2), East Buchan (Hamilton-Smith unpublished data); Nargun's Cave (NN1), Nowa Nowa 
(Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith 
unpublished data). 

Sphaeroceridae 

Leptocera sp., Tp, Gp. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith 
unpublished data). 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Paradox Cave (J48), Jenolan 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et 
al. 2003); VICTORIA: Wilson's Cave (EB4), East Buchan (Hamilton-Smith unpublished data); 
Nargun's Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Panmure Cave (H5), 
Mount Napier (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool 
(Hamilton-Smith unpublished data). 

Streblidae 

Ascodipteron archboldi Maa, Tp, Gx, P. QUEENSLAND: Chillagoe Caves (Maa 1971); Gordon 
Mine (Maa 1971). 

Ascodipteron australiense Muir, Tp, Gx, P. QUEENSLAND: Mount Etna Main Cave (El), Mount 
Etna (Maa 1971). 

Brachytarsina amboinensis uniformis Maa, Tp, Gx, P. NEW SOUTH WALES: Bungonia various 
caves (Maa 1971; Hamilton-Smith 1972; Eberhard 1998); Ashford Main Cave (AS1), Ashford (Maa 
1971); Back Creek Mine (Maa 1971); Belfery [sic] Cave (TR2), Timor (Maa 1971); Carrai Bat 
Cave (SC5), Stockyard Creek (Maa 1971); Drum Cave (B13), Bungonia (Maa 1971); Endless Cave 
(Maa 1971); Fig Tree Cave (W148), Wombeyan (Maa 1971); Prospect Tunnel (Maa 1971); Rise 
and Shine Mine (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); Yessabah Bat 
Cave (YE1), Yessabah (Maa 1971); QUEENSLAND: Chillagoe (Maa 1971); Viator Cave (VR4), 
Viator Hill, southern Queensland (Maa 1971); Mount Etna Main Cave (El), Mount Etna (Maa 
1971). 

Brachytarsina verecunda Maa, Tp, Gx, P. NEW SOUTH WALES: Ashford Main Cave (AS1), 
Ashford (Maa 1971); Bonalbo Colliery (Maa 1971); Cliefden (Maa 1971); Drum Cave (B13), 
Bungonia (Maa 1971); Humicrib Cave (WJ34), Wee Jasper (Maa 1971); Tanja Gold Mine (Maa 

Proc. Linn. Soc. N.S.W., 125, 2004 35 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 

1971); Morapabah Cave (Temagog Cave) (MP1), Morapabah (Maa 1971); Timor Caves (Maa 
1971); Wee Jasper (Maa 1971); Willi Willi Bat Cave (WW1), Willi Willi (Maa 1971); Bungonia 
various caves (Maa 1971; Hamilton-Smith 1972; Eberhard 1998); QUEENSLAND: Mount Etna 
Main Cave (El), Mount Etna (Maa 1971). 

Undetermined genus and species, Tp, Gx, P. QUEENSLAND: Riverton Main Cave (RN1), Riverton 
(Hamilton-Smith unpublished data); Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data). 

?Therevidae 

Undetermined genus and species, Gx?. NEW SOUTH WALES: Colong Main Cave (CGI), Colong 
(Hamilton-Smith unpublished data). 

Tipulidae 

Undetermined genus and species, Gx?. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan 
(Hamilton-Smith unpublished data). 

Trichoceridae 

Undetermined genus and species, Tp, Gp?. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Snowflake Cave (LI), 
Genelg River (Hamilton-Smith unpublished data); Fox Cave (U22), Naracoorte (Hamilton-Smith 
unpublished data); VICTORIA: Mount Widderin Cave (HI), Skipton (Hamilton-Smith unpublished 
data). 

Undetermined Family 

Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Cliefden Main Cave (CL1), 
Cliefden (Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton-Smith 
unpublished data); Tuglow Cave (Tl), Tuglow (Hamilton-Smith unpublished data); 
QUEENSLAND: Bat Cleft (E6), Mount Etna (Hamilton-Smith unpublished data); Speaking Tube 
(E7), Mount Etna (Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (Bl), 
Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); 
Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), 
Murrindal (Yen and Milledge 1990); Anticline Cave (Mil), Murrindal (Yen and Milledge 1990). 

Undetermined genus and species, Tp, Gp. VICTORIA: Bat Cave (P6), Portland (Hamilton-Smith 
unpublished data); Mt Widderin Cave (HI), Skipton (Hamilton-Smith unpublished data). 

Order Lepidoptera 

Noctuidae 

Agrotis infusa Boisduval, Tx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith 
unpublished data). 

Persectania ewingii Westwood, Tx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- 
Smith unpublished data). 

Pseudaletia australis Feaud., Tx. VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith 
unpublished data). 

Pyralidae 

Pyralis manihotalis Guenee, Tpl, Gp. QUEENSLAND: Rope Ladder Cave (FR2), Mingella, 
Fanning River (Weinstein and Edwards 1994). 

Pyralinae or Epipaschiinae sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) 
(Humphreys and Eberhard 2001). 



36 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 

Tineidae 

Lindera tessellatella Blanchard, Gb?. NEW SOUTH WALES: Humicrib Cave (WJ34), Wee Jasper 
(Eberhard and Spate 1995). 

Monopis crocicapitella Clemens, Tp, Gb. NEW SOUTH WALES: Drum Cave (B13), Bungonia 
(Eberhard 1998); Grill Cave (B44), Bungonia (Eberhard 1998); SOUTH AUSTRALIA: Bat Cave 
(U2), Naracoorte (Bellati et al. 2003); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- 
Smith unpublished data). 

Monopis sp., Gb. NEW SOUTH WALES: Gable Cave (CL7), Cliefden (Eberhard and Spate 1995); 
Colong Main Cave (CG3), Colong (Eberhard and Spate 1995); Jenolan undetermined cave (Gibian 
et al. 1988); Basin Cave (W4), Wombeyan (Smith 1982b); Undetermined caves, Wombeyan (Dew 
1963); Signature Cave (WJ7), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave 
(WJ8), Wee Jasper (Hamilton-Smith unpublished data); Dogleg Cave (WJ10), Wee Jasper 
(Eberhard 1993); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Humicrib 
Cave (WJ34), (Eberhard 1993); Carey's Cave (WJ100), Wee Jasper (Eberhard 1993); 
NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 
1971); Mullamullang Cave (N37) (Richards 1971); Cocklebiddy Cave (N48) (Richards 1971); 
Moonera Tank Cave (N53) (Richards 1971); Thylacine Hole (N63) (Richards 1971); Old 
Homestead Cave (N83) (Richards 1971); Dingo Cave (N160) (Richards 1971). 

Undetermined genus and species, Gb. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) 
(Humphreys and Eberhard 2001); Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 
2001); NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith 
unpublished data); Cliefden Main Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Willi 
Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); 
QUEENSLAND: Rope Ladder Cave (FR2), Mingella, Fanning River (Weinstein and Slaney 1995); 
Queenslander Tower (CH5246), Chillagoe (Matts 1987); Spring Tower (CH5223-5), Chillagoe 
(Marts 1987); Donna Tower (CH5155), Chillagoe (Matts 1987); Royal Arch Tower (CH5158-9), 
Chillagoe (Matts 1987); Tea Tree Tower (CH5137), Chillagoe (Matts 1987); Ryan Imperial Tower 
(CH5239), Chillagoe (Matts 1987); Wallaroo Tower (CH5201), Chillagoe (Matts 1987); Tower of 
London Cave (CH5) Chillagoe (Matts 1987); Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum 
Cave), Camooweal (Hamilton-Smith unpublished data); Holy Jump Lava Cave (BM1), Bauer's 
Mountain (Hamilton-Smith unpublished data); VICTORIA: Anticline Cave (Mil), Murrindal (Yen 
and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990); Nargun's Cave 
(NN1), Nowa Nowa (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool 
(Hamilton-Smith unpublished data). 

Undetermined Family 

Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) 
(Humphreys and Eberhard 2001); Swiflet Cave (CI30) (Humphreys and Eberhard 2001); Managers 
Alcove (CI50) (Humphreys and Eberhard 2001); Grimes Cave (CI53) (Humphreys and Eberhard 
2001); Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 2001). 

Undetermined genus and species, NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith 
unpublished data). 

Order Hymenoptera 

Braconidae 

Apanteles Icarpatus Say, Tpl, Gp. NEW SOUTH WALES: Humidicrib Cave (WJ34), Wee Jasper 
(Eberhard and Spate 1995); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); 
VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). 



Proc. Linn. Soc. N.S.W., 125, 2004 37 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Apanteles sp., Tp, Gp. NEW SOUTH WALES: Church Cave (W31), Wee Jasper (Hamilton-Smith 
unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith 
unpublished data). 

Undetermined genus and species. Tp?. QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer's 
Mountain (Hamilton-Smith unpublished data). 

Formicidae 

Amblyopone australis Erichson, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith 
unpublished data). 

Iridomyrmex purpureus Smith, Tx, Gx. SOUTH AUSTRALIA: Eregunda Mine near Blinman, 
Flinders Ranges (T. Moulds unpublished data). 

Oligomyrmex sp., Tp?, Gx?. QUEENSLAND: Crazy Cracks Cave, Jacks Gorge, Broken River (T. 
Moulds unpublished data). 

Pachycondyla sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) 
(Humphreys and Eberhard 2001). 

Undetermined genus and species, NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper 
(Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe 
(Hamilton-Smith unpublished data); Spring Cave, Mount Surprise (Hamilton-Smith unpublished 
data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Ichneumonidae 

Undetermined Cryptinae genus and species, Gp?. NEW SOUTH WALES: undetermined caves 
(Hamilton-Smith 1967); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith 
unpublished data); Spring Creek Cave (Bl), Buchan (Hamilton-Smith unpublished data); Wilson's 
Cave (EB4), East Buchan (Hamilton-Smith unpublished data). 

Myrmaridae 

Gonatocerinae sp., Gp?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 

Undetermined Family 

Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998). 

Undetermined genus and species, Gp?. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper 
(Hamilton-Smith unpublished data); Willi Willi Bat Cave (WW1), Willi Willi (Hamilton-Smith 
unpublished data); VICTORIA: Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished 
data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). 



38 Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



ACKNOWLEDGEMENTS 

This work was only possible due to the financial support 
of the Department of Environment and Heritage, South 
Australia, and the University of Adelaide. Thanks to Stefan 
Eberhard and Sue White for providing the stimulus for 
writing this paper. Many thanks to Elery Hamilton-Smith 
whose critical comments and unlimited access to his personal 
unpublished records greatly improved this paper. Thank you 
to Mike Gray, Courtenay Smithers and Judy Bellati who 
brought additional references to my attention. I also wish to 
thank John Jennings and Andy Austin for editorial comments. 



REFERENCES 

Beier, M. (1967). Some Pseudoscorpionidea from 

Australia, chiefly from caves. Australian 

Zoologist 14, 199-205. 
Beier, M. (1968). Some cave-dwelling 

Pseudoscorpionidea from Australia and New 

Caledonia. Records of the South Australian 

Museum 15, 757-765. 
Beier, M. (1975). Neue Pseudoskorpione aus Australien 

und Neu-Guinea. Annalen des Naturhistorischen 

Museums Wien 78, 203-213. 
Beier, M. (1976). A cavernicolous atemnid 

pseudoscorpion from New South Wales. 

Journal of the Australian Entomological Society 

15, 271-272. 
Bellati, J. (2001). Guanophilic invertebrate diversity and 

seasonality in Bat Cave at Naracoorte, South 

Australia. B. Sc. (Hons.). Department of 

Applied and Molecular Ecology, University of 

Adelaide, Adelaide. 
Bellati, J., Austin, A. D. and Stevens, N. B. (2003). 

Arthropod diversity of Bat Cave, Naracoorte 

Caves World Heritage Area, South Australia. 

Records of the South Australian Museum, 

Monograph Series 7, 257-265. 
Calder, D. R. and Bleakney, J. S. (1965). Microarthropod 

ecology of a porcupine-inhabited cave in Nova 

Scotia. Ecology 46, 895-899. 
Decu, V. (1986). Some considerations on the bat guano 

synusia. Travaux Institut de Speologie "Emile 

Racovitza" 25, 41-51. 
Dennis, T. P. (1986). Specimens of amphipod and isopod 

observed at Wombeyan Caves, NSW, Top End 

Speleological Society Inc. 
Dennis, T. P. and Mayhew, B. J. (1986). Specimens of 

Collembola and Psocoptera observed at 

Wombeyan Caves, NSW, Top End 

Speleological Society Inc. 
Dew, B. (1963). Animal life in caves. Australian Natural 

History 15, 158-161. 
Eberhard, S. M. (1993). Invertebrate cave fauna in NSW - 

Part 1, NSW Parks and Wildlife Service, 1-31. 
Eberhard, S. M. and Spate, A. (1995). Cave invertebrate 

survey, toward an atlas of NSW cave fauna. 



NSW heritage assistance program NEP 94765, 

1-112. 
Eberhard, S. M. (1998). Cave invertebrates. In 'Under 

Bungonia' . (Eds J. Bauer and P. Bauer). Pp. 74- 

83, 274-279. (Aiken Press, Sydney). 
Ferreira, R. L. and Martins, R. P. (1998). Diversity and 

distributions of spiders associated with bat 

guano piles in Mirrinho Cave (Bahia State, 

Brazil). Diversity and Distributions 4, 235-241. 
Ferreira, R. L. and Martins, R. P. (1999). Trophic 

structure and natural history of bat guano 

invertebrate communities, with special reference 

to Brazilian caves. Tropical Zoology 12, 231- 

252. 
Forster, R. R., Platnick, N. I. and Gray, M. R. (1987). A 

review of the spider superfamilies 

Hypochiloidea and Austrochiloidea (Areneae: 

Araneomorphae). Bulletin of the American 

Museum of Natural History 185, 1-116. 
Fricke, F. T. (1964). A note on Aulacopris reichi White 

(Col.: Scarabidae: Coprinae). Journal of the 

Entomological Society of Australia (NSW) 1, 36. 
Gibian, M., Smith, G. and Wheeler, L. (1988). Interim 

report on the survey of the invertebrate fauna of 

Jenolan Caves, Jenolan Caves Reserve Trust, 8. 
Gillieson, D. (1997). 'Caves: Processes, development and 

management'. (Blackwell: Oxford). 
Gnaspini, P. (1992). Bat guano ecosystems. A new 

classification and some considerations, with 

special references to Neotropical data. Memoires 

de Biospeologie 19, 135-138. 
Gnaspini, P. and Trajano, E. (2000). Chapter 13. Guano 

communities in tropical caves. In 'Ecosystems 

of the world. Subterranean ecosystems'. (Eds H. 

Wilkens, D. C. Culver and W. F. Humphreys). 

(Elsevier: Amsterdam). 
Gray, M. R. (1973a). Cavernicolous spiders from the 

Nullarbor Plain and south-east Australia. 

Journal of the Australian Entomological Society 

12, 207-221. 
Gray, M. R. (1973b). Survey of the spider fauna of 

Australian caves. Helictite 11, 47-75. 
Gray, M. R. (1992). New desid spiders (Araneae: 

Desidae) from New Caledonia and eastern 

Australia. Records of the Australian Museum 

44, 253-262. 
Gray, M. R. (1994). A review of filistatid spiders 

(Araneae: Filistatidae: Prithinae) from Australia 

with notes on subfamily classification. Records 

of the Australian Museum 46. 
Halliday, R. B. (1998). 'Mites of Australia. A checklist 

and bibliography'. (CSIRO Publishing: 

Melbourne). 
Halliday, R. B. (2000). The Australian species of 

Macrocheles (Acarina: Macrochelidae). 

Invertebrate Taxonomy 14, 273-326. 
Hamilton-Smith, E. (1966a). Pselaphidae from Australian 

caves. Journal of the Entomological Society of 

Queensland 5, 70-7 1 . 



Proc. Linn. Soc. N.S.W., 125, 2004 



39 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Hamilton-Smith, E. (1966b). Free-living Acarina in 
Australian caves. Bulletin of the National 
Speleological Society 28, 100-103. 

Hamilton-Smith, E. and Adams, D. J. H. (1966). The 

alleged obligate ectoparasitism of Myotyphlus 
jasoni (Matth.) (Coleoptera Staphylinidae). 
Journal of the Entomological Society of 
Queensland 5, 44-45. 

Hamilton-Smith, E. (1967). The Arthropoda of Australian 
caves. Journal of the Australian Entomological 
Society 6, 103-118. 

Hamilton-Smith, E. (1968). The insect fauna of Widderin 
Cave, Skipton Victoria. Victorian Naturalist 85, 
294-296. 

Hamilton-Smith, E. (1972). The bat population of the 
Naracoorte caves area. Proceedings of the 8th 
Conference of the Australian Speleological 
Society, Australian Speleological Society. 

Hamilton-Smith, E., Holland, E. G., Mott, K. and Spate, 
A. (1989). Cutta Cutta Caves Nature Park draft 
plan of management, Conservation Commission 
of the Northern Territory, 18-20. 

Hamilton-Smith, E. (2000). Report on current changes in 
biodiversity of the Bat Cave, Naracoorte World 
Heritage Area., Internal report for the 
Department of Environment and Heritage, South 
Australia. 

Harris, J. A. (1970). Bat guano cave environment. Science 
169, 1342-1343. 

Harris, J. A. (1971). Dynamics of a bat-guano cave 

ecosystem with particular reference to the guano 
mite Uroobovella coprophila (Womersley). PhD 
Thesis. University of Queensland, Brisbane. 

Harris, J. A. (1973). Structure and dynamics of a cave 
population of the guano mite, Uroobovella 
coprophila (Womersley). Australian Journal of 
Zoology 21, 239-275. 

Harvey, M. S. (1989). Two new cavernicolous chthoniids 
from Australia, with notes on the generic 
placement of the south-western Pacific species 
attributed to the genera Paraliochthonius Beier 
and Morikawia Chamberlain 
(Pseudoscorpionida: Chthoniidae). Bulletin of 
the British Arachnological Society 8, 21-29. 

Hoch, H. (1988). A new cavernicolous planthopper 
species (Homoptera: Fulgoroidea: Cixiidae) 
from Mexico. Mitteilungen der Schweizerischen 
entomologischen Gesellschaft 61, 295-302. 

Hoch, H. and Howarth, F. G. (1999). Multiple cave 

invasions by species of the planthopper genus 
Oliarus in Hawaii (Homoptera: Fulgoroidea: 
Cixiidae). Zoological Journal of the Linnean 
Society 111, 453-475. 

Horst, R. (1972). Bats as primary producers in an 
ecosystem. Bulletin of the National 
Speleological Society 34, 49-54. 

Howarth, F. G. (1983). Ecology of cave arthropods. 
Annual Review of Entomology 28, 365-389. 

Howarth, F. G. (1988). Environmental ecology of north 
Queensland caves: or why are there so many 



troglobites in Australia. Proceedings of the 1 7th 

biennial Australian Speleological Federation 

Tropicon Conference, Lake Tinaroo, far north 

Queensland., 76-83. 
Humphreys, W. F. (1991). Experimental re-establishment 

of pulse-driven populations in a terrestrial 

troglobite community. Journal of Animal 

Ecology 60: 609-623. 
Humphreys, W. F. and Eberhard, S. (2001). Subterranean 

fauna of Christmas Island, Indian Ocean. 

Helictite 37, 59-73. 
James, J. M., Middleton, G, Montgomery, N., Parker, F., 

Rolls, D., Scott, P. and Wellings, G (1976). 

'Timor Caves'. (Sydney Speleological Society: 

Sydney). 
Krantz, G W. and Fillipponi, A. (1964). Acari della 

famigilia Macrochelidae (Mesostigmata) nella 

collezione del South Australian Museum. 

Rivista di Parassitologia 25, 35-54. 
Lea, A. M. (1923). Australian dung beetles of the Sub- 
Family Coprides. Records of the South 

Australian Museum 2, 353-396. 
Lowry, J. W. J. (1996). Fauna found in Arramall River, 

Weelawadji and the Stockyard Gully Caves. 

http://wasg.iinet.net.au/eneabba.html. 
Maa, T. C. (1971). Studies in Batflies (Diptera: Streblidae: 

Nycteribiidae). Pacific Insects Monographs 28, 

1-247. 
Mackerras, M. J. (1965). Australian Blattidae (Blattodea) 

I. General remarks and revision of the Genus 

Polyzosteria Burmeister. Australian Journal of 

Zoology 13, 841-882. 
Mackerras, M. J. (1967). A blind cockroach from caves in 

the Nullarbor Plain (Blattodea: Blattellidae). 

Journal of the Australian Entomological Society 

6, 39-44. 
Martin, B. J. (1977). The influence of patterns of guano 

renewal on bat guano arthropod communities, 

Cave Reserve Foundation. Annual Report 1976, 

36-42. 
Mathews, E. G (1986). A revision of the troglophilic 

genus Brises Pascoe, with a discussion of the 

Cyphaleini (Coleoptera: Tenebrionidae). 

Records of the South Australian Museum 19, 77- 

90. 
Mathews, P. (1985). 'Australian Karst Index'. (Australian 

Speleological Federation Inc.: Sydney). 
Matts, G. (1987). History and summary of research in 

Chillagoe by Brother Nicholas Sullivan to end 

1986. Helictite 25, 74-78. 
Mc Alpine, D. K. (1966). Description and biology of an 

Australian species of Cypselosomatidae 

(Diptera) with a discussion of family 

relationships. Australian Journal of Zoology 14, 

673-685. 
Mc Alpine, D. K. (1993). A new genus of Australian 

cypselosomatid flies (Diptera: Nerioidea). 

General and Applied Entomology 25, 1-4. 
Medway, L. (1962). The swiftlets (Collocalia) of Niah 

Cave, Sarawak. Part 2 Ecology and the 

regulation of breeding. Ibis 104, 228-245. 



40 



Proc. Linn. Soc. N.S.W., 125, 2004 



T. MOULDS 



Moore, B. P. (1962). Notes on Australian Carabidae III. A 
remarkable cave-frequenting Harpaline from 
Western Victoria. Entomologist's Monthly 
Magazine 97, 188-190. 

Moore, B. P. (1964). New cavemicolous Carabidae 

(Coleoptera) from mainland Australia. Journal 
of the Entomological Society of Queensland 3, 
69-74. 

Moore, B. P. (1967). New Australian cave Carabidae 
(Coleoptera). Proceedings of the Linnean 
Society of New South Wales 91, 179-184. 

Moore, B. P., Humphreys, W. F., Decu, V. and Juberthie, 
C. (2001). Australie. In 'Encyclopaedia 
Biospeologica'. (Eds C. Juberthie and V. Decu). 
(International Society of Biospeleogy: Moulis, 
France). 

Moulds, T. A. (2003). Arthropod ecology of Bat Cave, 

Naracoorte, South Australia. Proceedings of the 
24th Biennial Conference of the Australian 
Speleological Federation, Bunbury WA, 
Australian Speleological Federation. 

Muchmore, W. B. (1982). A new cavemicolous 
Sathrochthonius from Australia 
(Pseudoscorpionida: Chthoniidae). Pacific 
Insects 24, 156-158. 

Park, O. and Barr, T. C. J. (1961). Some observations on a 
cave cricket (Abstr.). Bulletin of the 
Entomological Society of America 7, 144. 

Peck, S. B. (1976). The effect of cave entrances on the 
distribution of cave inhabiting terrestrial 
arthropods. International Journal of Speleology 
8, 309-321. 

Peck, S. B. and Christiansen, K. (1990). Evolution and 

zoogeography of the invertebrate cave faunas of 
the Driftless area of the Upper Mississippi River 
Valley of Iowa, Minnesota, Wisconsin, and 
Illinois, U.S.A. Canadian Journal of Zoology 
68, 73-88. 

Poulson, T. L. (1972). Bat guano ecosystems. Bulletin of 
the National Speleological Society 34, 55-59. 

Princis, K. (1963). Two new Western Australian 

cockroaches. Journal of the Royal Society of 
Western Australia 46, 11-12. 

Richards, A. M. (1964). The Rhaphidophoridae 

(Orthoptera) of Australia 2. A new genus. 
Proceedings of the Linnean Society of New 
South Wales 89, 373-379. 

Richards, A. M. (1966). The Rhaphidophoridae 

(Orthoptera) of Australia 3. A new genus from 
south-eastern Australia. Pacific Insects 8, 617- 
628. 

Richards, A. M. (1967a). Cockroaches (Blattodea) from 
Australian caves. Helictite 5, 35-44. 

Richards, A. M. (1967b). Abstract of Free living mites 
(Acarina) in Australian caves. Helictite 5, 70. 

Richards, A. M. (1971). An ecological study of the 
cavemicolous fauna of the Nullarbor Plain 
southern Australia. Journal of the Zoological 
Society of London 164, 1-60. 



Smith, G. (1982a). Cave fauna. In 'Wombeyan Caves'. 

(Eds H. J. Dyson, R. Ellis and J. M. James). 

173-178. (Sydney Speleological Society 

Occasional Paper) 
Smith, G. (1982b). Broad similarites in the invertebrate 

fauna in N.S.W. caves. Highland Caving Group 

Journal 3, 26-30. 
Smithers, C. N. (1964). New records of cave and mine- 
dwelling Psocoptera in Australia. Journal of the 

Entomological Society of Queensland 3, 85. 
Smithers, C. N. (1975). New Psocoptera records from 

Australian caves. Australian Entomological 

Magazine 2, 45-46. 
Snow, D. W. (1975). Oilbirds, cave-living birds of South 

America. Studies in Speleology 2, 257-264 +257 

plates. 
Trapani, J. (1997). Neotoma cinerea (Bushy-Tailed 

Woodrat, Packrat). http:// 

animaldiversity.ummz.umich.edu/accounts/ 

neotoma/n. cinerea.html. 
Vandel, A. (1973). Les Isopodes terrestres de 1' Australie. 

Etude systematique et biogeographique. 

Memoir es du Museum national d'Historie 

naturelle. Paris 82, 1-171. 
Waite, E. R. (1898). Note and exhibits. Proceedings of the 

Linnean Society of New South Wales 33, 803. 
Weinstein, P. and Edwards, E. D. (1994). Troglophilic 

moths in Australia: first record of a self- 
sustaining population. Journal of the Australian 

Entomological Society 33, 377-379. 
Weinstein, P. and Slaney, D. (1995). Invertebrate faunal 

survey of Rope Ladder Cave, Northern 

Queensland, a comparative study of sampling 

methods. Journal of the Australian 

Entomological Society 34, 233-236. 
Wellings, G. (1977). Invertebrate fauna of Bungonia 

caves. In 'Bungonia Caves'. (Eds R. Ellis, L. 

Hawkins, R. Hawkins, J. M. James, G. 

Middleton, B. Nurse and G. Wellings). Pp. 150- 
Williams, G. (2003). New distribution and biological 

records for native dung beetles, in the tribe 

Scarabaeini, from northern New South Wales. 

Proceedings of the Linnean Society of New 

South Wales 124, 13-16. 
Womersley, H. and Domrow, R. (1959). A new 

Asternolaelaps from Australia (Acarina: 

Ichthyostomatogasteridae). Records of the South 

Australian Museum 13, 355-358. 
Womersley, H. (1963a). Two new species of Acarina from 

guano from Australian bat caves. Transactions 

of the Royal Society of South Australia 86, 147- 

154. 
Womersley, H. (1963b). A new larval Neotrombidium 

(Acarina: Leeuwenhoekidae) from bat guano. 

Records of the South Australian Museum 14, 

473-476. 
Yen, A. L. and Milledge, G. A. (1990). Invertebrates of 

the Buchan-Murrindal area cave systems. 

Melbourne, Department of Conservation and 

Environment, Pp. 1-38. 



Proc. Linn. Soc. N.S.W., 125, 2004 



41 



CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 



Zumpt, F. and Patterson, P. M. (1952). Three new 
parasitic mites from the Ethiopian region 
(Acarina: Laelaptidae). Journal of the 
Entomological Society of Southern Africa 15, 
159-164. 



42 Proc. Linn. Soc. N.S.W., 125, 2004 



A Devonian Brachythoracid Arthrodire Skull (Placoderm Fish) 
from the Broken River Area, Queensland 

Gavin C. Young 

Department of Earth and Marine Sciences, Australian National University, Canberra ACT 0200 

gyoung@geology.anu.edu.au 

Young, G.C. (2004). A Devonian brachythoracid arthrodire skull (placoderm fish) from the Broken River 
area, Queensland. Proceedings of the Linnean Society of New South Wales 125, 43-56. 

An incomplete brachythoracid arthrodire skull acid-prepared from the Devonian limestones of the 
Broken River area of Queensland is described as Doseyosteus talenti gen. et sp. nov. It supposedly comes 
from strata dated by conodonts as late Early Devonian in age (Emsian stage), but shows several derived 
features of the skull, typical of Middle-Late Devonian brachythoracids, and not seen in any arthrodire from 
the Emsian limestones of the Burrinjuck area of NSW. The alignment with conodont zones of stratigraphic 
subdivisions of the Burrinjuck sequence is revised. Published information on the provenance and age of all 
previously described placoderm taxa from Broken River is reviewed and amended. The new taxon may be 
most closely related to Late Devonian (Frasnian) brachythoracids from Iran and the Gogo Formation of 
Western Australia. 

Manuscript received 27 May 2003, accepted for publication 22 Oct 2003. 

KEYWORDS: Placoderm fishes, Arthrodira, Brachythoraci, Broken River, Devonian, new genus 
Doseyosteus, Queensland. 



INTRODUCTION 

Devonian sedimentary rocks, including many 
marine limestones, are well exposed in the Broken 
River area of Queensland (Fig. 1). Conodonts form 
the basis for dating the sedimentary sequence (Mawson 
and Talent 1989; Sloan et al. 1995). Vertebrate remains 
reported from this sequence include microfossils from 
many horizons (De Pomeroy 1996; Turner, Basden 
and Burrow 2000), and less well known vertebrate 
macro-remains. The latter include two genera of 
antiarch placoderms described by Young (1990), a 
ptyctodont toothplate ascribed to IPtyctodus sp. by 
Turner and Cook (1997), a new species of the 
brachythoracid arthrodire Atlantidosteus Lelievre 1984 
described by Young (2003 a), an isolated suborbital 
plate of another arthrodire illustrated by Turner et al. 
(2000, fig. 8.7), and jaw remains of an onychodontid 
(Turner et al. 2000, fig. 5). Undescribed vertebrate 
macro-remains include various placoderm bones, most 
of which belong to brachythoracid arthrodires. The 
Arthrodira is the most diverse order within the class 
Placodermi, and its major subgroup, the Brachythoraci, 
comprises nearly 60% of about 170 genera within the 
Arthrodira (Carr 1995). The brachythoracid arthrodires 
were one of the most successful groups of early 



gnathostome fishes (e.g. Young 1986; Janvier 1996). 
In marine environments of the Late Devonian they 
included probably the largest predators of their time. 
The major radiation of brachythoracid subgroups had 
apparently already occurred by the Middle Devonian, 
and primitive representatives were already widespread 
in shallow marine environments of the Early Devonian 
(e.g. Young et al. 2001 ; Mark-Kurik and Young 2003), 
and are important in considering the origins and 
interrelationships of major brachythoracid subgroups 
(e.g. Lelievre 1995). 

The stratigraphic occurrence of various 
placoderm remains in the Broken River sequence were 
reviewed by Young (1993, 1995, 1996), De Pomeroy 
(1995, 1996), and Turner et al. (2000), and they have 
been mentioned in relation to conodont studies by 
Sloan et al. (1995). There has been conflicting 
information published about the provenance of some 
of the described placoderm taxa. These were collected 
from the Broken River area many years ago by 
Professor John Jell, University of Queensland, and sent 
to Canberra for acid preparation and study. In this paper 
I describe a new arthrodire skull from this collection, 
and review the locality information and age 
determinations for previously described placoderm 
taxa. 



DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 



unconformity 



CAINOZOIC [incl. basalt] 
L. DEVONIAN- 
CARBONIFEROUS 



- „~^\ MIDDLE DEVONIAN 
Tjf— gf jr [Broken River Group] 
disconformity 

i; i ;i; i ; i; i| early devonian 
tv'. *.t silurian 

unconformity 

7CAMBRIAN-ORDOVICIAN 
FOSSIL FISH LOCALITIES 




4 km 



Figure 1. (A) location of the Broken River area in Queensland, Australia; (B) geological 
map of the collecting area (modified from Turner, Basden and Burrow 2000, fig. 2), 
showing localities for previously described placoderm taxa, and the specimen described 
in this paper (ANU V1026). 



LOCALITY AND AGE OF DESCRIBED 

PLACODERM TAXA FROM THE BROKEN 

RIVER AREA 

Wurungulepis denisoni Young 1990 

According to information provided with this 
specimen, it came from University of Queensland 



locality L4399 (not L4339, given in error by Young 
1990: 45), on the north bank of the Broken River, Grid 
Reference 640 460 on the Burges 1:100 000 sheet, 
and was assigned a Middle Devonian (?Eifelian) age 
within the Broken River Formation (J.S. Jell, letter of 
17 April 1980). Judging by the map of the area 



44 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.C. YOUNG 



published by Sloan et al. (1995: fig. 2), the 
locality lies within outcrop referred to as 
'undifferentiated Broken River Group'. 

A ' Wurungulepis- Atlantidosteus 
fauna', of assumed Eifelian age, was listed in 
the macro vertebrate zonation of Young (1993, 
1995, 1996). However De Pomeroy (1995: 480) 
assigned Wurungulepis to the late Emsian 
serotinus Conodont Zone (CZ), citing a personal 
communication of J. A. Talent. This information 
was repeated by Turner et al. (2000: 498). Later 
(pers. comm. 28/8/95) J.A. Talent had advised 
A. Basden that this specimen was collected from 
the grid reference cited above, situated on a bend 
of the Broken River in an anticline, in strata 
which were pre-Dosey Limestone in the 
sequence, and equivalent to the Bracteata 
Formation and Lomandra Limestone (spanning 
the Emsian-Eifelian boundary; Sloan et al. 1995 : 
fig. 3). 

No conodont data were obtained from 
the specimen, so its precise position relative to 
the standard conodont zonation is uncertain. 
Wurungulepis is an early representative of the 
asterolepidoid antiarchs, with a high short trunk 
armour (Young 1990), and was placed within 
the asterolepidoid clade adjacent to 
Sherbonaspis, and as sister group to Stegolepis, 
Asterolepis, Remigolepis and Pambulaspis, by 
Zhu (1996: fig. 29). As earlier discussed (Young 
1990: 48) the initially suggested Eifelian age 
was consistent with the oldest asterolepid 
(pterichthyodid) occurrence in Europe, cited as 
Gerdalepis from the Eifelian of Germany by 
Denison (1978), although this occurrence is 
slightly younger (early Givetian) according to 
Otto (1998: 118). However Gardiner (1994) 
cited Young (1974) for an older record (Emsian) 
of the asterolepid antiarchs, but the 'cf. 
Pterichthyodes' mentioned by Young (1974) 
was based on an erroneous attribution by Hills 
(1958: 88) to the Early Devonian limestone 
sequence of an 'Antiarchan from Taemas'. In 
fact, the specimen in question came from the 
overlying Hatchery Creek Formation, of 
presumed Eifelian age (Fig. 2). This specimen 
was assigned to the new genus Sherbonaspis by 
Young and Gorter (1981). Previously, the 
suggested Emsian age of a pterichthyodid 
antiarch from the Georgina Basin (Young 
1984a) was noted as possibly the oldest 
occurrence of this group anywhere recorded. 
New evidence now indicates that two assemblages may 
have been mixed in this region (Burrow and Young, 



* 


C 






LU 


.2 






Q 


£ 


costatus 




■ 


LU 


partitus 


HC 






patulus 




serotinus 


? V1370 
I 









■6^=L C R| 


— 




C 


inversus- 


I 5 I 






.5 


laticostatus 


I4 






'<o 





R— 1 


— 


z 

< 


E 

LU 


perbonus- 


■1 3 




gronbergi 




z 

o 

> 






IB BJ 








leu cul 




LU 
Q 




dehiscens 


P Y SYJ 

|M M| 




>- 










-J 
< 






E B c m~ 

in < 


C 


pireneae 


LU 


(0 

'5> 

v. 


kindlei 


Q. " "s — 

2 E 




Q- 




LU 

LU 






sulcatus 


S 



Figure 2. Proposed alignment with conodont zones 
of subdivisions of the Early Devonian limestone 
sequence (Murrumbidgee Group) around 
Burrinjuck Dam, N.S.W., revised from Basden et 
al. (2000: fig. 2). Abbreviations for stratigraphic 
subdivisions are: B - Bloomfield Limestone 
Member; CB - Cavan Formation; CR - Crinoidal 
Limestone Member; CU - Currajong Limestone 
Member; HC - Hatchery Creek Formation; M - 
Majurgong Formation; R - Receptaculites 
Limestone Member; SY - Spirifer yassensis 
Limestone Member; W - Warroo Limestone 
Member; 1-6 - units of Upper Reef Formation. 
V1370 - horizon for highest known arthrodire in 
the sequence. 



in press), with the limestone occurrence yielding the 
antiarch probably younger than the diverse 



Proc. Linn. Soc. N.S.W., 125, 2004 



45 



DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 



Wuttagoonaspis fauna from underlying sandstones 
(Young and Goujet 2003). 

The antiarchs are a major subgroup of the 
class Placodermi, ranging in age from Early Silurian 
to latest Devonian. In recent years there has been a 
significant expansion in our knowledge of the group. 
A cladistic analysis of their distribution in relation to 
phylogeny by Young (1984b) involved 22 taxa and 
40 characters. In a recent review of antiarch phylogeny, 
Zhu (1996) noted some 45 genera and 154 species, 
and his data matrix used 66 characters for 40 genera. 
The original age assessment of Eifelian for 
Wurungulepis from Broken River is most consistent 
with our current knowledge of this large and diverse 
group. 

Nawagiaspis wadeae Young 1990 

This specimen is recorded from locality 
BRJ68D (University of Queensland locality L4428; 
'small limestone outcrop on eastern side of gully 1 
km upstream from Six Mile yard'), Grid Reference 
596 442 on the Burges 1:100 000 sheet, which was 
assigned a Middle Devonian (?Givetian) age within 
the Broken River Formation (J.S. Jell, letter of 17 April 
1980). Apparently this specimen was found by Dr 
Mary Wade. 

Again, De Pomeroy (1995: 480) referred this 
taxon to the significantly older (late Emsian) serotinus 
CZ, based on its assigned position within the Bracteata 
Formation in section Br4 of Sloan et al. (1995, fig. 6). 
This information was repeated by Turner et al. (2000: 
498, 506). However Prof. J. A. Talent's previous advice 
to the author (pers. comm. 5/8/92), was that this 
specimen was considerably younger (ensensis - varcus 
Zones; late Eifelian - Givetian). Clearly, there was 
some confusion about which fish specimen was being 
referred to. Subsequent advice given to A. Basden 
(pers. comm. 28/8/95), was that N. wadeae came from 
the bank of Dosey Creek (Grid Reference 616 437), 
the location of section Br2 within outcrop of the 
Bracteata Formation (Sloan et al. 1995: fig. 2). The 
different, and presumably correct, locality information 
provided with the specimen, as cited above, 
corresponds to the vicinity of the boundary between 
the Papilio and Mytton Formations on the map of Sloan 
et al. (1995: fig. 2). This is consistent with the Givetian 
age first suggested by J.S. Jell. 

Nawagiaspis wadeae is another antiarch, 
originally interpreted as possibly a primitive 
bothriolepidoid (Young 1990), although in Zhu's 
(1996) phylogeny it comes out as a basal 
asterolepidoid. Apart from primitive Chinese antiarchs, 
and the erroneous Emsian pterichthyodid occurrence 
discussed above, the stratigraphic record of this group 



is Middle-Late Devonian (Gardiner 1994, fig. 32.1). 
The bothriolepidoid clade had an earlier history in Asia, 
and apparently expanded its range to most regions of 
the world in the Givetian (Young 2003b). 

The confusion about the provenance of this 
specimen may have resulted from the misconception 
that it was a recognisable 'skull' when collected. 
Turner et al. (2000) used this term to refer to the type, 
but the specimen as collected was a largely complete 
trunk armour, and the incomplete skull, missing its 
central portion, formed a minor part of the specimen. 
The whole specimen may have appeared to a non- 
vertebrate worker to represent a 'skull'. Such fish 
remains, when collected in the field, are generally not 
determinable until after acid preparation (e.g. the type 
specimen of Atlantidosteus pacifica Young 2003a, 
before preparation, was assumed to be a ventral plate 
of the trunk armour, rather than a large suborbital bone 
from the cheek). 

A summary list of prepared fish remains from 
the original J.S. Jell collection was provided to J.A. 
Talent in 1995 to check on age and locality data. This 
list mentioned only one skull, the brachythoracid 
specimen described below, of which locality data 
provided by J.S. Jell are almost the same as stated by 
Sloan et al. (1995) for N. wadeae. Thus it seems that 
the specimen described below, previously listed as a 
'skull', has been confused with the type of N. wadeae, 
leading to erroneous locality and age information being 
given in De Pomeroy (1995), Sloan et al. (1995), and 
Turner et al. (2000). In the context of the global 
distribution in time and space of this major placoderm 
subgroup (see above), it is almost certain that 
Nawagiaspis is Middle Devonian in age, and a Givetian 
age, as first suggested by J.S. Jell, is most consistent 
with other information about the stratigraphic 
distribution of the more derived antiarchs. 

Atlantidosteus pacifica Young 2003a 

This specimen came from locality BRJ 67B 
(University of Queensland locality L 4472), Grid 
Reference 675 485 on the Burges 1:100 000 sheet, 
described as 'Top of ridge to three-quarters way down 
western slope, west of road between Six Mile Dam 
and Diggers Creek' (J.S. Jell, letter of 17 April 1980). 
This is the locality (with a slightly different grid 
reference) referred to as 'Fish Hill' by Turner et al. 
(2000: 507). They assigned it a middle Eifelian age 
(costatus - australis conodont zone), but noted that 
Sloan et al. (1995) gave a slightly longer partitus - 
early kockelianus zonal range for the Fish Hill section. 
This is consistent with the original assignment of a 
Middle Devonian (?Eifelian) age within the Broken 
River Formation by Prof. J.S. Jell. This occurrence is 



46 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.C. YOUNG 



part of the evidence for proposing an Eifelian 
'Wurungulepis-Atlantidosteus fauna' in the 
macro vertebrate zonation of Young (1993, 1995, 
1996). 

Doseyosteus talenti gen. et sp. nov. 

This specimen, described below, was the only 
one in the J.S. Jell collection lacking a sample number 
at the time of preparation. It is highly probable that it 
was a sample collected the year before the other 
material, and was taken to Canberra separately by Dr 
P. Jell (J.S. Jell, letter of 17 April 1980). The following 
locality details, provided by Prof. J.S. Jell (letter of 17 
April 1980), indicate that it is the specimen collected 
from the alternative erroneous locality for Nawagiaspis 
just discussed: 'BRJ34 = L 4054. Grid Reference 616 
438 Burges 1:100,000 sheet. Western bank of Dosey 
Creek, 750 m upstream from its junction with Broken 
River. Base of thick limestone lens in Broken River 
Formation, Middle Devonian. ? Eifelian' . 

In a published listing of University of 
Queensland locality numbers (Turner et al. 2000: 506), 
UQL4054 is assigned to 'basal part of limestone, 
Lomandra/Dosey Limestone, Broken River Group', 
with a slightly different grid reference (615 438), but 
the same locality description as above. However, it is 
assigned to the Emsian serotinus CZ, citing Sloan et 
al. (1995). 

Again, no conodonts were obtained from the 
sample, and section Br4 through the Bracteata 
Formation at this locality did not produce identifiable 
conodonts (Sloan et al. 1995: caption to fig. 6). 
Nevertheless, these authors (p.5) considered the entire 
formation to belong to the serotinus CZ, making it 
equivalent to the upper part of the Burrinjuck (NSW) 
limestone sequence, which extends from the top of 
the pirenae CZ (latest Pragian) into the serotinus CZ 
(the second youngest zone of the late Emsian). It is 
therefore relevant to make comparisons with the 
stratigraphic distribution of the diverse arthrodire 
assemblage described from the Burrinjuck limestone 
sequence. 

The described arthrodire fauna from the 
Burrinjuck sequence (White 1952, 1978; White and 
Toombs 1972; Young 1979, 1981, in press a, b; Young 
et al. 2001; Mark-Kurik and Young 2003) includes 10 
genera of brachythoracids, amongst which the most 
derived taxa (Cathlesichthys and Dhanguura) come 
from the upper part of the Wee Jasper limestone 
sequence. Basden et al. (2000, fig. 2) showed the 
youngest arthrodire skull from the Wee Jasper section 
(ANU VI 370; the holotype of Dhanguura) to come 
from the uppermost unit 6 of the 'Upper Reef 
Formation' of Young (1969). This specimen is more 



advanced than other arthrodires known from the 
Burrinjuck sequence in possessing several derived 
characters of the skull, the most obvious being the T- 
shaped rostral plate, a feature of more derived 
eubrachythoracids (character 5 of Carr 1991; character 
4 of Lelievre 1995). Eubrachythoracids were the most 
diverse fish group of the Middle and Late Devonian, 
and the new Broken River brachythoracid described 
below clearly belongs to this group, with a skull which 
is more advanced in several respects than any of the 
known Burrinjuck arthrodires (see below). Gardiner 
(1994) lists the first occurrence of this grouping (his 
family Coccosteidae) as Coccosteus Miller 1841 from 
the Middle Devonian (Eifelian) of Scotland, for which 
a late Eifelian age is indicated by spores of the 
devonicus-naumovae zone (V.T. Young 1995). The 
same species (Coccosteus cuspidatus) is recorded from 
the Kernave Member of the Narva Formation in the 
Baltic sequence, although a related brachythoracid 
'Protitanichthys' occurs a little earlier, and in 
equivalent strata (costatus CZ) in the Rhenish sequence 
(Mark-Kurik 2000). However Otto (1997: 115) 
suggested that remains of early eubrachythoracids 
(coccosteids) first occur in the early Eifelian of 
Scotland, Germany, and the Baltic sequence. 

Dhanguura johnstoni Young (in press a) 
comes from a horizon about 420 m stratigraphically 
above the boundary equivalent of the Bloomfield and 
Receptaculites Members of the Taemas Limestone. A 
similar horizon high in the limestone sequence has 
produced the large lungfish Dipnorhynchus cathlesae 
Campbell and Barwick 1999. The lungfish locality is 
close to localities L537 and L538 of Pedder et al. 
(1970) which yielded tetracorals Vepresiphyllum 
dumosum, Sulcorphyllum pavimentum, 

Chalcidophyllum vesper and C. gigas. This represents 
the uppermost 'tetracoral teilzone' of the 
Murrumbidgee Group (Pedder et al. 1970: fig. 4), and 
is Coral Fauna F in the scheme of Garratt and Wright 
(1989). These authors considered the succeeding G 
and H Coral Faunas to overlap, and belong to the late 
Emsian, rather than Eifelian as previously assessed. 
Garratt and Wright (1989) also aligned Coral Fauna F 
from Wee Jasper (and the Sulcor Limestone of northern 
NSW) with the mid-Emsian inversus CZ (see column 
13 of Young 1995, 1996). However Basden et al. 
(2000: fig. 2) showed the uppermost beds of the 
limestone sequence at Wee Jasper (containing Coral 
Fauna F) extending well into the next youngest 
serotinus CZ. Evidence supporting this (summarised 
by Basden 2001, table 2.1) derives from reassignment 
of some of the conodonts from the highest productive 
sample (C62) in Pedder et al.'s (1970) section 2, 
referred by them to Polygnathus linguiformis 



Proc. Linn. Soc. N.S.W., 125, 2004 



47 



DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 



linguiformis, but reassigned to Polygnathus inversus 
by Klapper and Johnson (1975), and to Polygnathus 
serotinus (delta morphotype) by Mawson (1987). On 
the other hand, the age in terms of conodont zone 
alignment of several constituent members of the 
Taemas Limestone, as indicated by Basden et al. (2000, 
fig. 2), seem to be too young, and should be revised 
downwards on the following evidence. Lindley (2002a: 
275) noted that the occurrence of the index species of 
Coral Fauna D (Chalcidophyllum recessum) in the 
Currajong Limestone Member indicates that it should 
be aligned with the dehiscens rather than the perbonus 
CZ. The overlying Bloomfield Limestone Member 
may also have lower beds of dehiscens rather than the 
perbonus CZ age (Basden 2001 : table 2. 1). The Warroo 
Limestone Member contains perbonus CZ elements 
(Nicoll, in Lindley 2002b), and the uppermost 
Crinoidal Limestone Member in the Taemas sequence 
may align with both the inversus and the serotinus CZ 
(Basden 2001: table 2.1). 

These revised alignments are summarised in 
Fig. 2. Correlation with the upper part of the Wee 
Jasper sequence is unclear, because the constituent 
members of the Taemas Limestone are difficult to 
recognise in the thicker upper part of the sequence, 
represented by units 1-6 of Young (1969). If the new 
arthrodire skull described below from Broken River 
is of serotinus CZ age, as proposed by Sloan et al. 
(1995), it is still considerably more derived (see below) 
than any arthrodire from the Burrinjuck sequence. If 
correctly dated, this would indicate that derived 
features characterising the Middle-Late Devonian 
eubrachythoracid arthrodires had originated at least 
by late Emsian time. 

To summarise, it is emphasised that there is 
no overlap in the arthrodire skull characters just 
discussed between the Burrinjuck and Broken River 
limestone sequences, even though the youngest 
occurrences in the former sequence are also the most 
derived taxa within the better-documented Burrinjuck 
arthrodire fauna. For the new taxon described below, 
this evidence would support either a latest Emsian age 
(but younger than the Burrinjuck sequence), or an 
Eifelian age as originally suggested by Prof. J.S. Jell. 



anth, anterior nuchal thickening; 

Ce, central plate; 

cf.Ce, area overlapping Ce plate; 

cf.M, area overlapping marginal plate; 

cf.PM, area overlapping postmarginal plate; 

cf.PtO, area overlapping postorbital plate; 

cr.im, inframarginal crista; 

esc, central sensory line canal; 

d.end, openings of dermal tube for endolymphatic 

duct; 

dep, depression; 

gr.M, groove on Ce plate which received the edge of 

the marginal plate; 

ifc.ot, otic branch of infraorbital sensory groove; 

if.r, infranuchal ridge; 

if.pt, infranuchal pit; 

kb, knob-like thickening of inframarginal crista; 

lcp, lateral consolidated part of skull roof; 

lie, main lateral line sensory canal; 

M, marginal plate; 

mp, middle pitline; 

mppr, posterior median process of nuchal plate; 

Nu, nuchal plate; 

oa.Ce, area overlapped by Ce plate; 

oa.M, area overlapped by M plate; 

oa.Nu, area overlapped by Nu plate; 

orb, orbital notch; 

Pi, pineal plate; 

plpr, posterolateral process or lobe on Ce plate; 

PM, postmarginal plate; 

pmc, postmarginal sensory groove; 

pnp, postnuchal process of paranuchal plate; 

PNu, paranuchal plate; 

pp, posterior pitline; 

PrO, preorbital plate; 

PtO, postorbital plate; 

R, rostral plate; 

soa, subobstantic area; 

soc, supraorbital sensory canal; 

th.end, endolymphatic thickening; 

th.pre, pre-endolymphatic thickening; 

tnth, transverse nuchal ridge or thickening; 

vg, vascular grooves. 



ABBREVIATIONS 



SYSTEMATIC PALAEONTOLOGY 



The specimen described below (prefix ANU 
V) is housed in the Earth and Marine Sciences 
Department, Australian National University, Canberra 
(GCY Vertebrate Collection). Standard abbreviations 
for placoderm dermal bones are used in the text and 
figures, and together with other morphological 
abbreviations are listed as follows: 



Class PLACODERMI McCoy, 1848 
Order ARTHRODIRA Woodward, 1891 
Suborder BRACHYTHORACI Gross, 1932 

Doseyosteus talenti gen. et sp. nov. 
Name 

From Dosey Creek, the type locality, and the 
Greek osteus (bone). The species name recognises 



48 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.C. YOUNG 



Professor John A. Talent, Macquarie University, who 
has had a long and distinguished career in Devonian 
research, including extensive work in the Broken River 
area of Queensland. 

Diagnosis 

A eubrachythoracid arthrodire in which the 
skull shows an embayed anterior margin of the nuchal 
plate resulting from overlap by the central plates, the 
central plates have strong posterolateral lobes 
separating the nuchal and paranuchal, and a mesial 
process of the marginal plate extends to the anterior 
angle of the paranuchal. Subobstantic area of skull 
extending onto marginal plate. Dermal bones smooth, 
or ornamented with fine tubercles. 

Remarks 

Since only the skull is known, and it is 
incomplete, several features characterising the derived 
subgroup 'Eubrachythoraci' are for the present inferred 
for this new taxon. Definition of the eubrachythoracid 
arthrodires is discussed by Carr (1991: 379-381) and 
Long (1995: 55). Thus Doseyosteus talenti gen. et sp. 
nov. is assumed to have had a T-shaped rostral plate, a 
posteriorly placed pineal plate separating the 
preorbitals, a dermal process of the preorbital plate 
forming the anterodorsal margin of the orbit, and 
trilobate central plates. The holotype shows a strongly 
developed posterior thickening of the skull roof, which 
in the midline is represented by the anterior nuchal 
thickening. This is much more prominent than the 
transverse ridge on the posterior margin of the nuchal 
plate, and is a derived feature seen in coccosteomorph 
and pachyosteomorph brachythoracids, but generally 
lacking in Early Devonian taxa, for example the genus 
Cathlesichthys from Burrinjuck, NSW (Young, in press 
a). The embayed anterior margin and inferred 
proportions of the nuchal plate, and the strong 
posterolateral lobe of the Ce plate, are resemblances 
to the Late Devonian taxa Eastmanosteus and 
Golshanichthys, but the former differs in having the 
posterior pitline well developed on the posterolateral 
lobe of the central plate, and both forms lack the mesial 
process of the marginal plate inferred for this new 
taxon. 

Material 

ANU VI 026 (holotype), an incomplete skull 
preserved as two unconnected portions. 

Locality and Horizon 

Locality BRJ34 (University of Queensland 
locality L4054), Grid Reference 616 438, Burges 1 : 100 
000 sheet; western bank of Dosey Creek, 750 m 



upstream from its junction with Broken River (J.S. Jell, 
letter of 17 April 1980; see discussion above). Horizon 
was described as the 'base of thick limestone lens in 
Broken River Formation', assigned to the Bracteata 
Formation (Sloan etal. 1995) or the 'Lomandra/Dosey 
Limestone, Broken River Group (Turner et al. 2000). 
Age: ?late Emsian - Eifelian (see discussion above). 

Description 

ANU V1026 represents a large part of the 
posterolateral region of a brachythoracid skull roof, 
preserved as two separate portions. The larger portion 
(Fig. 3 A,D) includes parts of the Nu, PNu and Ce plates 
(Fig. 4A,B), and the right postmarginal corner of the 
skull is preserved as a separate portion (Figs. 3B,C, 
4C,D). The specimen was extracted from the rock in 
six pieces, but they are well preserved, suggesting that 
it was broken up before incorporation in the sediment. 
The nuchal (Nu) plate is represented by most of its 
right half, including the midline, so its overall shape 
can be estimated. Midline length of the Nu is about 70 
mm. It has an embayed posterior margin, with a 
prominent posterior median process (mppr, Fig. 4). 
Except for the posterior lateral corner the right lateral 
margin of the Nu plate is fairly well displayed on the 
external surface. The bone is fractured in its middle 
region, and shows anteriorly that it was both 
overlapped and underlapped by the central (Ce) plate, 
a condition also reported in Holonema (Miles 1971). 
Along the anterior margin of the plate a thin 
overlapping lamina of the Ce plate has broken away 
to reveal an extensive overlap area (oa.Ce, Fig. 4B). 
In unbroken condition the anterior margin of the Nu 
plate would have been deeply embayed (Fig. 5). On 
its visceral surface extensive contact faces for the 
central plates are developed in the normal manner 
(cf.Ce, Fig. 4A). Other features shown are the 
prominent infranuchal pits (if.pt) and ridge (if.r) and 
the transverse nuchal thickening or ridge (tnfh). 

Noteworthy is the strong development of the 
anterior nuchal thickening (anth), which is relevant to 
the question of the age of this specimen (see discussion 
above). This is a derived feature of brachythoracids, 
and in ANU VI 026 is more pronounced than in any 
Emsian brachythoracid from the Burrinjuck fauna. 
These have Nu plates which are fairly flat in front of 
the infranuchal pits. This is the case even in a form 
like Cathlesichthys, which is derived in having a very 
strong transverse nuchal ridge (Young in press a). In 
posterior view ANU VI 026 shows that the anterior 
nuchal thickening is more pronounced than the 
transverse nuchal ridge, the reverse of the condition 
in Cathlesichthys. This advanced character is also seen 
in most Middle-Late Devonian brachythoracids, such 



Proc. Linn. Soc. N.S.W., 125, 2004 



49 



DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 




Figure 3. Doseyosteus talenti gen. et sp. nov. Holotype (ANU V1026). Larger (A,D) and smaller (B,C) 
skull portions in external (B,D) and internal (A,C) views. 



as Golshanichthys, Tafilalichthys, and various Gogo 
forms (e.g. Lelievre et al. 1981; Lelievre 1991; Miles 
and Dennis 1979; Long 1988, 1995; Dennis-Bryan 
1987). These taxa all resemble the giant Famennian 
form Dunkleosteus, where the 'posterior consolidated 
arch' of the skull roof ('PCA' of Heintz 1932: fig. 13) 
is a broad thickening running in front of the infranuchal 
pits, as the main transverse thickening of the skull. In 
contrast, in the Early Devonian form Cathie sichthys 
from Burrinjuck, the transverse nuchal ridge located 
behind the infranuchal pits forms the main thickening 
supporting the posterior skull margin. 

The right paranuchal (PNu) plate of 
Doseyosteus gen. nov. is represented externally by an 
elongate portion including the mesial margin forming 
sutures with the Nu and Ce plates (PNu, Fig. 4B). There 
is also a small broken part of the postnuchal process 
(pnp). The PNu and Ce plates were also connected by 
a complex interlocking suture; a broken part around 



the anterior end of the PNu exposes an overlap area 
(oa.Ce, Fig. 4B), and the edge of a more extensive 
contact face is shown on the visceral surface (cf.Ce, 
Fig. 4A). The endolymphatic thickening forms a broad 
thickened area mesially (th.end), combining with the 
thickened portion of the Nu plate (anth). This thickened 
part of the skull is much more prominent than in 
primitive brachythoracids like Buchanosteus or 
Taemasosteus (White 1978; Young 1979). Along the 
broken edge of the specimen, maximum bone thickness 
(in the part enclosing the endolymphatic duct) is almost 
15 mm, which is three times the bone thickness at the 
anterior preserved extremity of the Nu. The exoskeletal 
division of the right endolymphatic duct opens on the 
visceral skull roof surface at the anterior edge of the 
area of thickened bone (th.end), and is also visible on 
the broken margin of the specimen (d.end, 
Fig. 4A). This is also an advanced character of the 
brachythoracid skull - in large Emsian brachythoracids 



50 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.C. YOUNG 



cf.M 



?oa.M 




esc 



\M-m&-i*' - -^j/ 


*lcp 


ifc. 


\ M 

Ot\ j 


r^Sk \A 






/** cr.im 
cf.PM 






pmc' N 


X&s^- soa 



Figure 4. Doseyosteus talenti gen. et sp. nov. Holotype (ANU V1026). A,B- Larger portion of skull in 
internal (A) and external (B) views. C,D. Smaller skull portion in internal (C) and external (D) views. 



from Burrinjuck the endolymphatic duct is not within 
the bone, but anteriorly forms a bony tube attached to 
or projecting from the inner surface of a much thinner 
PNu plate (Young in press a: figs. 3, 4, 7A, 9B). A 
similar condition occurs in Holonema from Gogo (JA. 
Long, pers. comm.; Miles 1971: fig. 53). 

The preserved part of the right Ce plate is 
crossed by a prominent sensory groove (esc), which 



must be the central sensory canal rather than the 
supraorbital sensory canal, because of its oblique 
orientation to the midline. Middle and posterior pitlines 
are represented by faint markings in the region of the 
ossification centre (mp, pp). Anterolateral and 
posterolateral margins of the preserved part of the Ce 
plate are somewhat fractured, but appear to 
approximate natural margins. The former is bevelled 



Proc. Linn. Soc. N.S.W., 125, 2004 



51 



DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 




x \ ! 10 mm 



Figure 5. Doseyosteus talenti gen. et sp. nov. Attempted skull roof reconstruction, preserved 
portions shaded. 



externally, and internally shows a contact face for the 
postorbital plate (cf.PtO), showing that it overlapped 
the PtO extensively, as in most other brachythoracids 
(e.g. Miles and Westoll 1968: fig. 2; Young 1979: fig. 
1; 1981: fig. 5). Holonema is an exception in this 
respect (Miles 1971: fig. 12). Subdivisions of the 
posterior part of this contact face suggest that it also 
overlapped the marginal (M) plate (cf.M, Fig. 4A). 

The posterolateral margin of the Ce plate is 
somewhat thicker, and carries a deep groove (gr.M) 
for an interlocking suture, the Ce plate providing 
external and internal laminae to enclose the margin of 
the contiguous bone. The nature of the preserved 
margins suggests that they approximate the suture 



position. Since the anterior end of the PNu is well 
shown on the specimen, and is most unlikely to have 
extended to this margin of the Ce plate, it seems that 
the intervening space must have been occupied by a 
mesial projection of the M plate (M, Fig. 5). This 
arrangement has not previously been recorded in 
brachythoracids. A similar but smaller process of the 
M intrudes the Ce plate of Buchanosteus, but this is 
some distance in front of the PNu (Young 1979: fig. 

1). 

There is a long posterolateral projection of 
the Ce plate partly separating the Nu and PNu plates 
(plpr, Fig. 4B), a feature seen in various other 
brachythoracids. An early example with this 



52 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.C. YOUNG 



morphology is Ulrichosteus Lelievre, 1982a from the 
Givetian of Germany, but this form has the Nu plate 
extending anteriorly in front of the PNu, whereas in 
Doseyosteus the PNu is slightly longer. Ardennosteus 
Lelievre, 1982b also has a strong posterolateral lobe 
of the Ce, but this Famennian form differs in its sinuous 
interlocking sutures, broader transverse nuchal 
thickening, and coarse tubercular ornament. 
Development of a posterolateral lobe of the Ce is one 
of three features representing the 'trilobate' condition 
of the Ce plates (characters 13, 14, 21 of Carr 1991), a 
widespread condition amongst Middle-Late Devonian 
eubrachythoracids which has proved difficult to define. 
Internally this part of the Ce is more extensive, the 
overlapped portion extending back to the 
endolymphatic thickening, again as in other 
brachythoracids. The visceral surface of the Ce is 
gently concave laterally, with several shallow grooves 
(vg) resembling the vascular grooves described in 
Holonema by Miles (1971: fig. 12). This depressed 
region is flanked mesially by the pre-endolymphatic 
thickening (th.pre), which forms a low broad ridge with 
a curved anteromesial orientation. The preserved 
anteromesial edge of the Ce plate is thickened and 
abraded (Fig. 3D). 

Associated with this skull portion was a 
smaller part of the left preobstantic corner of the skull 
roof (Fig. 3B,C), assumed to have belonged to the same 
individual. The specimen includes part of the PNu and 
M plates (Fig. 4C,D), and is crossed by a section of 
the main lateral line (He), and the infraorbital (if cot) 
and postmarginal (pmc) sensory canals. Unlike forms 
such as Coccosteus, Holonema and Buchanosteus 
(Miles and Westoll 1968; Miles 1971; Young 1979), 
the M plate carries part of the subobstantic area (soa, 
Fig. 4D). A subobstantic area of similar extent is seen 
in the Gogo brachythoracid Harrytoombsia Miles and 
Dennis (1979: fig. 4), and in all plourdosteids sensu 
Long (1995). The PM plate is missing, but on the 
visceral surface there is a clear contact face for this 
bone (cf.PM, Fig. 4C). The visceral surface also shows 
the inframarginal crista to be strongly developed, 
dorsally as a very prominent irregular knob of bone 
(kb) separated posteriorly by a deep groove from the 
ventrally directed crista (cr.im), which itself carries a 
groove. The free ventral margin of the plate is 
thickened (lep), representing the 'lateral consolidated 
part' of the skull, and a depression between the 
thickening and the inframarginal crista (dep) may 
correspond to similar structures in Coccosteus and 
Buchanosteus Young (1979: 314). 

The external ornament on both specimens 
comprises fine tubercles in some areas, sometimes only 
faintly discernible on a generally smooth surface (Fig. 



3B,D). The fine ornament is similar to that on the SO 
plate of Atlantidosteus pacifica, but that form displays 
affinity with the homostiid arthrodires in a range of 
features (Young 2003a), whereas the skull of 
Doseyosteus talenti gen. et sp. nov. lacks various 
specialised characters of Homostius and related forms 
(e.g. elongate Nu and PNu plates, small dorsal orbits, 
etc.). The reduced ornament also distinguishes this new 
form from various 'coccosteomorph' arthrodire 
remains known from the early Middle Devonian of 
northern Germany and the Baltic sequence (Otto 1997, 
1999). 

An attempted reconstruction of the skull roof 
of the new taxon based on available information is 
presented in Fig. 5. The skull could have been broader 
across the preobstantic corners than shown, since the 
gap between the two preserved portions is based only 
on a general alignment of sutures and sensory grooves. 
The anterior part of the skull is unknown, and restored 
shape of bones is generally based on various 
coccosteomorph arthrodires (e.g. Denison 1978: fig. 
57). Advanced features depicted (T-shaped R plate, 
Pi plate separating PrO plates, trilobate Ce plates) are 
based on their co-occurrence with preserved skull 
characters in all other known taxa. They need to be 
confirmed with additional material. On the larger 
preserved portion, the breadth and anterior embayment 
of the Nu plate, and the marked posterior lobe of the 
Ce plate separating the Nu and PNu plates, are general 
resemblances to Eastmanosteus and Golshanichthys, 
as noted above. The M and Ce plates retain extensive 
contact to separate the PNu from the PtO, the assumed 
primitive condition for brachythoracids. In contrast, 
the plourdosteid arthrodires, which were widespread 
in the Late Devonian, and apparently replaced the 
largely Middle Devonian coccosteids (Long 1995), 
have a much enlarged PtO reaching back to contact 
both the Ce and PNu plates. In consequence the M 
plate is reduced in size, whereas in Doseyosteus gen. 
nov., although not completely preserved, the M plate 
was clearly a more extensive bone, which apparently 
shows a unique feature in the large mesial process 
embaying the Ce plate in front of the PNu. 

In summary, this new but poorly known 
brachythoracid shows a range of advanced characters 
otherwise only seen in Middle or Late Devonian taxa, 
and it resembles the Frasnian taxa Eastmanosteus and 
Golshanichthys in several features which might 
indicate a close relationship. Eastmanosteus 
yunnanensis Wang, 1991 from the Givetian of China 
would otherwise be the earliest known member of this 
group (family Dinichthyidae). Kiangyousteus Liu, 
1955, also from China (Givetian of Szechuan), may 
be another primitive dinichthyid (Denison 1978). Both 



Proc. Linn. Soc. N.S.W., 125, 2004 



53 



DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 



taxa differ from the new form described here in their 
well-developed coarse tubercular ornament, 
presumably a primitive feature. Doseyosteus talenti 
gen. et sp. nov. displays an unusual shape of the M 
plate which is apparently unique to this new genus 
and species. More material, including the unknown 
trunk armour, which in brachythoracids comprises 17 
separate bones, will clarify the affinities of this new 
taxon. 



ACKNOWLEDGMENTS 

Professor J.S. Jell (University of Queensland) and 
Professor K.S.W. Campbell (ANU) are thanked for making 
the specimen available for study. Mr R.W. Brown 
(Geoscience Australia) assisted in acid preparation. Professor 
J.A. Talent and Dr A. Basden (Macquarie University) advised 
and discussed at length the provenance and age of Broken 
River placoderms, and Dr S. Turner (Queensland Museum) 
provided comparative material. Comparison with European 
and Moroccan arthrodire material was facilitated by a visiting 
professorship at the Museum national d'Histoire naturelle, 
Paris, in 1999. Professor D. Goujet is thanked for arranging 
this, and for the provision of facilities, and together with Dr. 
H. Lelievre and Dr. P. Janvier discussed at length placoderm 
morphology and relationships. Dr Lelievre arranged for 
arthrodire casts to be sent to Canberra for comparative study. 
B. Harrold is thanked for providing essential computer 
support at ANU, and V. Elder for assistance with specimen 
curation. Dr E. Mark-Kurik and Dr R. Carr discussed 
arthrodire phylogeny, and Dr Carr arranged for a visit to 
Cleveland, Ohio, for study of large arthrodire material. 
Financial support was provided in Canberra by ANU 
Faculties Research Fund Grants F01083 and F02059, and 
overseas by the Alexander von Humboldt Foundation, for a 
Humboldt Award in Berlin (2000-2001), and assistance with 
travel to the USA (Flagstaff and Cleveland, 2000). I thank 
Prof. H.-P. Schultze for provision of facilities in the Museum 
fiir Naturkunde, Berlin. Dr P. De Deckker is thanked for 
provision of facilities in the Geology Dept., ANU. This 
research was a contribution to IGCP Projects 328, 406, 410, 
and 491. 



REFERENCES 

Basden, A. (2001). 'Early Devonian fish faunas of eastern 
Australia: documentation and correlation'. Ph.D 
Thesis, Macquarie University [unpublished], 
349 pp., 63 figs. 

Basden, A., Burrow, C.J., Hocking, M., Parkes, R. and 
Young G.C. (2000). Siluro-Devonian 
microvertebrates from south-eastern Australia. 
In IGCP 328, Final Report, A. Blieck and S. 
Turner (eds), Courier Forschungsinstitut 
Senckenberg 223, 201-222. 

Burrow, C.J. and Young, G.C. (in press). Acanthodian 
fishes from the Cravens Peak Beds, Western 



Queensland. Memoirs of the Queensland 

Museum 
Campbell, K.S.W. and Barwick, R.E. (1999). A new 

species of the Devonian lungfish Dipnorhynchus 

from Wee Jasper, New South Wales. Records of 

the Australian Museum 51, 123-140. 
Carr, R.K. (1991). Reanalysis of Heintzichthys gouldii 

(Newberry), an aspinothoracid arthrodire 

(Placodermi) from the Famennian of northern 

Ohio, with a review of brachythoracid 

systematics. Zoological Journal of the Linnean 

Society 103, 349-390. 
Carr, R.K. (1995). Placoderm diversity and evolution. 

Bulletin du Museum national d'Histoire 

naturelle, Paris, Section C, 17, 85-125. 
De Pomeroy, A.M. (1995). Australian Devonian fish 

biostratigraphy in relation to conodont zonation. 

Courier Forschungsinstitut Senckenberg 182, 

475-486. 
De Pomeroy, A.M. (1996). Biostratigraphy of Early and 

Middle Devonian microvertebrates from Broken 

River, north Queensland. Records of the Western 

Australian Museum 17, 417-437. 
Denison, R.H. (1978). 'Placodermi. Handbook of 

Paleoichthyology, Volume 2'. (Ed. H.P. 

Schultze). (Gustav Fischer Verlag, Stuttgart, New 

York), 128 pp. 
Dennis-Bryan, K.D. 1987. A new species of eastmanosteid 

arthrodire (Pisces: Placodermi) from Gogo, 

Western Australia. Zoological Journal of the 

Linnean Society 90, 1-64. 
Gardiner, B.G. (1994). Placodermi. In 'The Fossil Record 

2' (Ed. M.J. Benton) pp. 583-588 (Chapman and 

Hall, London). 
Garratt, M.J. and Wright, A.J. 1989. Late Silurian to Early 

Devonian biostratigraphy of southeastern 

Australia. In 'Devonian of the World' (Eds. 

Mcmillan, N.J., Embry, A.F. and Glass, D.J.) 

Canadian Society of Petroleum Geologists, 

Calgary, Memoir 14(3), 647-662. 
Gross, W. (1932). Die Arthrodira Wildungens. 

Geologische u. Paldontologische Abhandlungen 

19, 5-61. 
Heintz, A. (1932). The structure of Dinichthys: a 

contribution to our knowledge of the Arthrodira. 

Bashford Dean Memorial Volume- Archaeic 

Fishes 4, 115-224. 
Hills, E.S. (1958). A brief review of Australian fossil 

vertebrates. In 'Studies on Fossil Vertebrates 

(Ed. T.S. Westoll) pp. 86-107 (Athlone Press, 

London). 
Janvier, P. (1996). 'Early Vertebrates'. (Clarendon Press, 

Oxford), 393 pp. 
Klapper, G. and Johnson, D.B. (1975). Sequence in the 

conodont genus Polygnathus in Lower 

Devonian at Lone Mountain, Nevada. Geologica 

et Palaeontologica 9, 65-77. 
Lelievre, H. (1982a). Ulrichosteus milesi n. g. n. sp. A new 

brachythoracid arthrodire (Placodermi) from the 

Givetian of the Rhineland. Neues Jahrbuch fur 

Geologie und Paldontologie Monatshefte 1982 



54 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.C. YOUNG 



(8), 501-508. 

Lelievre, H. (1982b). Ardennosteus ubaghsi n.g., n.sp., 

Brachythoraci primitif (vertebre, placoderme) du 
Famennien d'Esneux (Belgique). Annates de la 
Societe Geologique de Belgique 105, 1-7. 

Lelievre, H. (1984). Atlantidosteus hollardi n.g., n.sp., 

nouveau Brachythoraci (vertebres, placodermes) 
du Devonien inferieur du Maroc presaharien. 
Bulletin du Museum national d'Histoire 
naturelle, Paris 6, 197-208. 

Lelievre, H. (1991). New information on the structure and 
the systematic position of Tafilalichthys lavocati 
(placoderm, arthrodire) from the Late Devonian 
of Tafilalt, Morocco. In 'Early vertebrates and 
related problems of evolutionary biology' (Eds. 
M.M. Chang, Y.H. Liu, G.R. Zhang) pp. 121-130 
(Science Press, Beijing). 

Lelievre, H. (1995). Description of Maideria falipoui n.g., 
n. sp., a long snouted brachythoracid 
(Vertebrata, Placodermi, Arthrodira) from the 
Givetian of Maider (South Morocco), with a 
phylogenetic analysis of primitive 
brachythoracids. Bulletin du Museum national 
d'Histoire naturelle, Paris 17, 163-207. 

Lelievre, H., Janvier, P. and Goujet, D. (1981). Les 
vertebres Devonien de l'lran central: rV, 
arthrodires et ptyctodontes. Geobios 14, 677-709. 

Lindley, I.D. (2002a). Lower Devonian ischnacanthid fish 
(Gnathostomata: Acanthodii) from the Taemas 
Limestone, Lake Burrinjuck, New South Wales. 
Alcheringa 25, 269-291. 

Lindley, I.D. (2002b). Acanthodian, onychodontid and 

osteolepidid fish from the middle-upper Taemas 
Limestone (Early Devonian), Lake Burrinjuck, 
New South Wales. Alcheringa 26, 103-126. 

Liu, H.T. (1955). Kiangyousteus, a new arthrodiran fish 
from Szechuan, China. Acta Palaeontologica 
Sinica 3, 261-274. 

Long, J.A. (1988). A new camuropiscid arthrodire (Pisces: 
Placodermi) from Gogo, Western Australia. 
Zoological Journal of the Linnean Society 94, 
233-258. 

Long, J.A. (1995). A new plourdosteid arthrodire from the 
Late Devonian Gogo Formation, Western 
Australia: systematics and phylogenetic 
implications. Palaeontology 38, 39-65. 

Mark-Kurik, E. (2000). The Middle Devonian fishes of 
the Baltic States (Estonia, Latvia) and Belarus. 
Courier Forschungsinstitut Senckenberg, 223, 
309-324. 

Mark-Kurik, E. and Young, G.C. (2003). A new 

buchanosteid arthrodire (placoderm fish) from 
the Early Devonian of the Ural Mountains. 
Journal of Vertebrate Paleontology 23, 13-27. 

Mawson, R. (1987). Early Devonian conodont faunas 
from Buchan and Bindi, Victoria, Australia. 
Palaeontology 30, 251-297. 

Mawson, R. and Talent, J.A. (1989). Late Emsian- 

Givetian stratigraphy and conodont biofacies - 
carbonate slope and offshore shoal to sheltered 



lagoon and nearshore carbonate ramp - Broken 
River, north Queensland, Australia. Courier 
Forschungsinstitut Senckenberg, 117, 205-259. 

McCoy, F. (1848). On some new fossil fishes of the 

Carboniferous period. Annals and Magazine of 
Natural History 2, 1-10. 

Miles, R. S. (1971). The Holonematidae (placoderm 
fishes), a review based on new specimens of 
Holonema from the Upper Devonian of Western 
Australia. Philosophical Transactions of the 
Royal Society of London. B. Biological Sciences 
263, 101-234. 

Miles, R.S. and Dennis, K. (1979). A primitive 

eubrachythoracid arthrodire from Gogo, Western 
Australia. Zoological Journal of the Linnean 
Society 66, 31-62. 

Miles, R.S. and Westoll, T. S. (1968). The placoderm fish 
Coccosteus cuspidatus Miller ex Agassiz from the 
Middle Old Red Sandstone of Scotland. Part I. 
Descriptive morphology. Transactions of the 
Royal Society of Edinburgh 67, 373-476. 

Miller, H. (1841). 'The Old Red Sandstone'. (Johnstone 
and Hunter, Edinburgh). 

Otto, M. (1997). Vertebrate fossils of the Middle 

Devonian (Eifelian) Muhlenberg Formation in 
the Bergisches Land, northwestern Germany. 
Palaontologische Zeitschrift 71, 107-116. 

Otto, M. (1998). New finds of vertebrates in the Middle 
Devonian Brandenberg Group (Sauerland, 
Northwest Germany). Palaontologische 
Zeitschrift 12, 117-134. 

Otto, M. (1999). Neues Material von Protitanichthys? 

montanus (Vertebrata, Placodermi, Arthrodira) 
aus dem unteren Mitteldevon des Bergischen 
Landes. Neues Jahrbuch fur Geologie und 
Palaontologie Monatshefte 1999 (7), 397-408. 

Pedder, A.E.H., Jackson, J.H. and Philip, G.M. (1970). 
Lower Devonian biostratigraphy in the Wee 
Jasper region of New South Wales. Journal of 
Paleontology 44, 206-251. 

Sloan, T.R., Talent, J.A., Mawson, R., Simpson, A.J., 
Brock, G.A., Engelbretsen, M.J., Jell, J.S., 
Aung, A.K., Pfaffenritter, C, Trotter, J. and 
Withnall, I.W. (1995). Conodont data from 
Silurian-Middle Devonian carbonate fans, debris 
flows, allochthonous blocks and adjacent 
autochthonous platform margins: Broken River 
and Camel Creek areas, north Queensland, 
Australia. Courier Forschungsinstitut 
Senckenberg 182, 1-77. 

Turner, S., Basden, A. and Burrow, C.J. (2000). Devonian 
vertebrates of Queensland. In 'IGCP 328, Final 
Report' (Eds A. Blieck and S. Turner). Courier 
Forschungsinstitut Senckenberg 223, 487-521. 

Turner, S. and Cook, A. (1997). Ptyctodont jaw from the 
Broken River Province, NEQ. Memoirs of the 
Queensland Museum 42, 80. 

Wang J.-Q. (1991). A fossil Arthrodira from Panxi, 

Yunnan. Vertebrata PalAsiatica 29, 264-275 
(Chinese, English summary). 



Proc. Linn. Soc. N.S.W., 125, 2004 



55 



DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 



White, E.I. (1952). Australian Arthrodires. Bulletin of the 
British Museum (Natural History) (Geology), 1, 
249-304. 

White, E.I. (1978). The larger arthrodiran fishes from the 
area of the Burrinjuck Dam, N.S.W. 
Transactions of the Zoological Society of 
London 34, 149-262. 

White, E.I. and Toombs, H.A. (1972). The buchanosteid 
arthrodires of Australia. Bulletin of the British 
Museum (Natural History) (Geology) 22, 
379-419. 

Woodward, A.S. (1891). 'Catalogue of Fossil Fishes. Part 
2'. (British Museum (Natural History), London). 

Young, G.C. (1969). 'The geology of the Burrinjuck- Wee 
Jasper area, New South Wales'. B.Sc. Hons 
thesis, ANU Canberra [unpublished]. 

Young, G.C. (1974). Stratigraphic occurrence of some 
placoderm fishes in the Middle and Late 
Devonian. Newsletters on Stratigraphy 3, 243- 
261. 

Young, G.C. (1979). New information on the structure 

and relationships of Buchanosteus (Placodermi, 
Euarthrodira) from the Early Devonian of New 
South Wales. Zoological Journal of the Linnean 
Society 66, 309-352. 

Young, G.C. (1981). New Early Devonian 

brachythoracids (placoderm fishes) from the 
Taemas - Wee Jasper region of New South 
Wales. Alcheringa 5, 247-271. 

Young, G.C. (1984a). An asterolepidoid antiarch 

(placoderm fish) from the Early Devonian of the 
Georgina Basin, central Australia. Alcheringa 8, 
65-80. 

Young, G.C. (1984b). Comments on the phylogeny and 

biogeography of antiarchs (Devonian placoderm 
fishes), and the use of fossils in biogeography. 
Proceedings of the Linnean Society of New 
South Wales 107, 443-473. 

Young, G. C. (1986). The relationships of placoderm 
fishes. Zoological Journal of the Linnean 
Society 88, 1-57. 

Young, G.C. (1990). New antiarchs (Devonian placoderm 
fishes) from Queensland, with comments on 
placoderm phylogeny and biogeography. 
Memoirs of the Queensland Museum 28, 35-50. 

Young, G.C. (1993). Middle Palaeozoic macrovertebrate 
biostratigraphy of Eastern Gondwana. In 
'Palaeozoic Vertebrate Biostratigraphy and 
Biogeography'. (Ed. J. A. Long) pp. 208-251. 
(Belhaven Press, London). 

Young, G.C. (1995). Timescales 4. Devonian. 

Biostratigraphic charts and explanatory notes. 
2 nd Series. Australian Geological Survey 
Organisation, Record 1995/33, 1-47. 

Young, G.C. (1996). 'Devonian (chart 4)', In 'An 

Australian Phanerozoic Timescale' (Eds. Young, 
G.C. and Laurie, J.R.) pp. 96-109. (Oxford 
University Press, Melbourne). 

Young G.C. (2003a). A new species of Atlantidosteus 
Lelievre, 1984 (Placodermi, Arthrodira, 
Brachythoraci) from the Middle Devonian of the 



Broken River area (Queensland, Australia). 
Geodiversitas 25, 681-694. 

Young, G.C. (2003b). North Gondwanan mid-Palaeozoic 
connections with Euramerica and Asia; 
Devonian vertebrate evidence. Courier 
Forschungsinstitut Senckenberg 242, 169-185. 

Young, G.C. (in press a). Large brachythoracid arthrodires 
(placoderm fishes) from the Early Devonian of 
Wee Jasper, New South Wales, Australia. 
Journal of Vertebrate Paleontology 

Young, G.C. (in press b). Homostiid remains (placoderm 
fishes; Arthrodira), from the Early Devonian of 
the Burrinjuck area, New South Wales. 
Alcheringa 

Young, G.C. and Goiter, J.D. (1981). A new fish fauna of 
Middle Devonian age from the Taemas/Wee 
Jasper region of New South Wales. Bureau of 
Mineral Resources Geology & Geophysics, 
Bulletin 209, 83-147. 

Young, G.C. and Goujet, D. (2003). Devonian fish 

remains from the Dulcie Sandstone and Cravens 
Peak Beds, Georgina Basin, central Australia. 
Records of the Western Australian Museum, 
Supplement 65, 1-85. 

Young G.C, Lelievre H. and Goujet D. (2001). Primitive 
jaw structure in an articulated brachythoracid 
arthrodire (placoderm fish; Early Devonian) 
from southeastern Australia. Journal of 
Vertebrate Palaeontology 21, 670-678. 

Young, V.T. (1995). Micro-remains from Early and 

Middle Devonian acanthodian fishes from the 
U.K. and their biostratigraphic possibilities. 
Ichthyolith Issues, Special Publication 1, 65-68. 

Zhu, M. (1996). The phylogeny of the Antiarcha 

(Placodermi, Pisces), with the description of 
Early Devonian antiarchs from Qujing, Yunnan, 
China. Bulletin du Museum national d'Histoire 
naturelle, Paris (4,C) 18, 233-347. 



56 



Proc. Linn. Soc. N.S.W., 125, 2004 



Effects of Slashing and Burning on Thesium australe R. Brown 
(Santalaceae) in Coastal Grasslands of NSW 

Janet S. Cohn 

Biodiversity Research and Management Division, NSW National Parks and Wildlife Service, PO Box 1967, 

Hurstville, NSW 2220 (janet.cohn@npws.nsw.gov.au) 



Cohn, J. (2004). Effects of slashing and burning on Thesium australe R. Brown (Santalaceae) in coastal 
grasslands of NSW. Proceedings of the Linnean Society of New South Wales 125, 57-65. 

Two studies examined the effects of burning and cutting on aspects of the population dynamics of 
a nationally vulnerable herb, Thesium australe on the central and north coast of NSW. Study sites were 
grasslands dominated by Themeda australis with scattered native shrubs (Banksia integrifolia, Acacia 
sophorae) and the exotic shrub Chrysanthemoides monilifera ssp. rotundata. In the first study (May 1995 to 
December 1996), Thesium australe occurred at high density (1/m 2 ) on exposed, long-unburnt headlands. In 
the second study, (December 1996 to December 1998), Thesium australe was at low density (<l/100m 2 ) on 
more protected and recently burnt hinterland. On the headlands, winter treatments had no significant effect 
on the survival, density and vigour of Thesium australe. In the hinterland, one year after summer treatments, 
seedling recruitment resulted in a higher density of Thesium australe in the cut plots than either the burnt or 
the control. Flowering and fruiting of Thesium australe were not restricted by season. After winter and 
summer treatments, flowering and fruiting occurred within 6 months and 1 year, respectively. Although 
exposed coastal headlands may require no management intervention to increase the occurrence of Thesium 
australe, except where the possibility of shrub invasion exists, a regime of slashing on less exposed hinterlands 
may be needed to reduce competition from Themeda australis. Further research is necessary to determine if 
slashing or burning the more protected hinterland would yield different results if carried out in seasons other 
than summer. 

Manuscript received 1 March 2003, accepted for publication 22 October 2003. 

KEYWORDS: fire, grasslands, headlands, mowing, slashing, Thesium australe. 



INTRODUCTION Non-coastal, long-unburnt grasslands 

dominated by Themeda australis I triandra, have been 

Although Thesium australe is a herb with a shown to be s P ecies P oor < Stuwe and Parsons 1971 > 

wide ecological tolerance, extending from tropical to Kirkpatrick 1986; McDougall 1989), largely as aresult 

alpine climates, it is confined to widely scattered of the w g h competitive ability of this tussock grass 

locations in open woodlands and grasslands where ( Groves 1974 >- Wlth a § eneral recent declme m fire 

Themeda australis/ T. triandra (Kangaroo Grass) is frequency on coastal headlands (Griffith 1992), 

common in the understorey (Scarlett et al. 1994). On dominance by Themeda australis and the recruitment 

the north coast of NSW, T. australe occurs on grassy of native md exotic shrubs ** P otential threats t0 the 

headlands used predominantly for passive recreation, survival of T austmle ( Gnfflth 1992 )' * lthou S h Coo P er 

often adjacent to residential areas (Griffith 1992; Fig. < 1986 ) suggested that headlands exposed to salt-laden 

n winds may be an exception. He cites the persistence 

In south-eastern Australia, open woodland and of T australe at Perpendicular Point, 20 years after 

grassland communities have largely been modified and fire > as an exam P le - 

fragmented by introduced grazers, cultivation and Research on Thesium alpinum in Denmark 

changed fire regimes (Stuwe and Parsons 1977; Scarlett found that ll became extinct as a result of shadm § from 

and Parsons 1990; McDougall and Kirkpatrick 1994; ttees (Lojtnant and Worsoe 1980). Thesium australe 

Tremont and Mclntyre 1994; Prober and Thiele 1995; ma y be similarly sensitive. In coastal Victoria, an 

Lunt 1997). As a consequence T. australe is rated as increase in natlve shrub md *"* recruitment has been 

nationally vulnerable (Briggs and Leigh 1996) and hnked to a declme in fire frequency (Bennett 1994; 

vulnerable in NSW under Schedule 2 of the NSW McMahon et al. 1994; Lunt 1998a b). On the north 

Threatened Species Conservation Act 1995 coast of NSW ' increased recruitment of native shrubs 



EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 




Figure 1. Grassland habitat of Thesium australe 
at Look at Me Now Headland on the north coast 
of NSW. 



and trees {Acacia, Banksia, Allocasuarina spp.) and 
an invasive exotic shrub, Bitou bush, 
(Chrysanthemoid.es monolifera ssp. rotundata have 
been observed (Dodkin and Gilmore 1985; Griffith 
1987; Griffith 1992). 

A number of studies have suggested a regime 
of regular burning and/or mowing to maintain species 
richness in grasslands and prevent shrub invasion 
(Groves 1974; Stuwe and Parsons 1977; Kirkpatrick 
1986; McDougall 1989; Lunt 1990a, 1998b). Current 
information on the response of T. australe to fire in 
the field has been based on observations. While Leigh 
and Briggs (1989) suggest that survival and recruitment 
are unaffected by fire, Archer (1984) believed seeds 
were stimulated to germinate. In laboratory trials, 
Scarlett (pers. comm.) found that heat did not stimulate 
seed germination. There have been no studies on the 
effect of mowing or cutting on T. australe (Griffith 
1992). 

On coastal headlands and conservation reserves 
where burning or slashing grasslands may be used for 
conservation or hazard reduction purposes, it is 
important to establish their effect on native species. 
Two separate studies examined the effects of a single 
burning and a single cutting on aspects of the 
population dynamics of T. australe, namely: 
1/ its survival, density, vigour and reproductive status 

where it occurred at relatively high density on 

long-unburnt, exposed headlands (winter 

treatments); 
2/ its density and reproductive status where it was at 

very low density in a more protected and 

recently burnt hinterland (summer treatments). 

These were not intended to be comparative 
studies and indeed the different timing and methods 
of treatment (see Materials and Methods), driven by 



the availability of resources, make this not possible 
anyway. 

MATERIALS AND METHODS 

The studies were located at several sites on the 
north and central coast of NSW (Fig. 2): Perpendicular 
Point (AMGR Easting 485600, Northing 6499200); 
Look at Me Now Headland (E 518000, N 6661300); 
and Old Bar Park (E 461300, N 6462800). 
Perpendicular Point and Look at Me Now Headland 
are within respectively, Kattang Nature Reserve (NR) 
and Moonee Beach NR. Both are managed by the New 
South Wales National Parks and Wildlife Service 
(NSW NPWS). Old Bar Park is managed by The 
Greater Taree City Council. 

All three sites are used for recreation, 
predominantly by walkers. Although no motor 
vehicular access is allowed in the NRs, there was 
evidence of their past usage at Look at Me Now 
Headland, where at the time of this study wheel ruts 
were still very obvious. Vehicles were used on 
Perpendicular Point as recently as 1986 (Cooper 1986). 
There is some use of motor vehicles in Old Bar Park, 
but this is mostly on the pre-existing tracks and the 
airstrip (author's personal observations). 

Perpendicular Point and Look at Me Now 
Headland are characterised by black headland soils, 
which are loamy soils high in organic matter (Parbery 
1947). Yellow podzolic soils predominate at Old Bar 
Park (Long 1996). Aspects and slopes of the study sites 
varied. At Perpendicular Point the site was located on 
a north-western aspect with a slope of 9°, whilst at 
Look at Me Now Headland the site was on a more 
exposed southerly aspect with a slope of 6°. The site at 
Old Bar Park was flat. 

The study sites were in grassland communities 
dominated by Themeda australis. Scattered shrubs at 
Perpendicular Point included native (e.g. Acacia 
sophorae, Banksia integrifolia) and exotic (e.g. 
Chrysanthemoides monilifera ssp. rotundata) taxa. 
Another nationally endangered herb, Zieria prostrata 
(Briggs and Leigh 1996) also occurred on a number 
of the headlands with T. australe (Griffith 1992; NPWS 
1998). 

Thesium australe was found at Perpendicular 
Point in 1957 (Cooper 1986) and at Look at Me Now 
Headland and Old Bar Park after 1992 (Griffith 1992). 
Although at relatively high density at Perpendicular 
Point and Look at Me Now Headland (approximately 
1/m 2 ), at Old Bar Park it occurred mostly as very 
scattered plants (approximately <1/100 m 2 ). Thus, the 
focus at this latter site was more on recruitment 



58 



Proc. Linn. Soc. N.S.W., 125, 2004 



J.S.COHN 




Moonee Beach 
Nature Reserve 

Coffs Harbour 



50 



100 



kilometres 



Port Macquarie 



Kattang Nature 
Reserve 



Old Bar Park 




Figure 2. Locality of study sites within Moonee Beach NR (Look at Me Now Headland), Kattang NR 
(Perpendicular Point) and Old Bar Park on the north and central coast of NSW. Stippled areas represent 
estate managed by NSW National Parks and Wildlife Service. 



responses to treatments. 

There was little information on the fire history 
at the three sites. Griffith (1992) believed that 
Perpendicular Point may not have burnt for a 
considerable period of time. In 1985, Cooper (1986) 
believed that Perpendicular Point had not burnt for at 
least 20 years. There was no record of the last fire at 
Look at Me Now Headland. Old Bar Park was last 
burnt in 1991 by a low intensity fire (T. Cross pers. 
comm.). Approximately 1 year prior to this study Old 
Bar Park was slashed (S. Griffith pers. comm.), 
presumably for hazard reduction purposes. 

Headlands (high density plants) 

Treatments were applied in winter (July 1995) 
at Perpendicular Point and Look at Me Now Headland 
(Table 1). There were 15 replicate plots of each 
treatment (burnt, cut, control). Each treatment was 
allocated randomly to a 0.5 m x 0.5 m plot laid out in 
rows, over a total area of 75 m 2 at Perpendicular Point 
and 1 12 m 2 at Look at Me Now Headland. Plots were 



burnt using a gas burner. Because of the heavy dew, 
each burnt plot was subjected to heat for 5 minutes, 
until all of the grasses and herbs had been burnt and 
the bare ground had been heated and scorched. This 
simulated a high intensity burn (R. Bradstock pers. 
comm.). In the cutting treatment all grasses and herbs, 
including T. australe were cut to within 0.5 cm of the 
ground with shears. 

At both sites, in all plots, individual T. australe 
plants were tagged and numbered and the fates of the 
original and emerged plants were surveyed over 1.5 
years (Table 1 ). Data on plant vigour (number of stems/ 
plant; Perpendicular Point only) and the incidence of 
flowering or fruiting were also collected. 

Analyses of the proportion of T. australe plants 
surviving 6 and 16 months after treatment, were made 
using Generalised Linear Modelling (GLIM), with a 
binomial error structure (Crawley 1993). The effects 
of the factors, treatment (burnt, cut, control) and site 
(Perpendicular Point, Look at Me Now Headland) and 
their interactions were examined using the chi-squared 



Proc. Linn. Soc. N.S.W., 125, 2004 



59 



EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 



Table 1. The dates of treatment applications and monitoring at the study sites. 



Study Site 



Treatment (date) 



Monitoring Dates 

(pre and post treatment) 



Perpendicular Point 

Look at Me Now Headland 

Old Bar Reserve 



burn, cut (26/7/95) 
bum, cut (27/7/95) 
bum, slash (16/12/96) 



12/5/95, 14/2/96, 4/12/96 
26/7/95, 11/2/96,18/12/96 
3/12/96, 2/12/97, 16/12/98 



statistic. 

The density of T. australe plants (0.25 nr 2 ) was 
examined using fully factorial analyses of variance 
(ANOVA) and Tukey tests for pairwise comparisons. 
The effects of treatments (burnt, cut, control) and sites 
(Perpendicular Point, Look at Me Now Headland) were 
examined at pre- and post-treatment dates (0, 6 and 
16 months). To satisfy Cochran's test of homogeneity 
of variances, data were square root transformed and if 
necessary a more conservative level of significance 
(p<0.01) was applied (Underwood 1981). 

Analyses of the vigour of T. australe plants 
(number of stems/plant) at Perpendicular Point were 
made using one-way ANOVAs. The effects of 
treatments (burnt, cut, control) were examined at pre- 
and post- treatment dates (0, 6 and 16 months). Data 
from all plots and cohorts within each treatment were 
pooled. 

Hinterland (low density plants) 

At Old Bar Park, where T. australe occurred at 
very low density, large plots were subjected to burning 
or slashing. Each treatment (burnt, slashed, control) 
was allocated to a lOmx 10m plot within an overall 
area of 40 m x 50 m. There were 2 replicates of each 
treatment. Whilst for practical purposes the two burnt 
plots were placed together, replicates of the cut 
treatment and control were randomly allocated to the 
remaining plots. Burning took place in hot conditions 
during summer (December 1996). Two plots were 
slashed the next day to within 5 cm of the ground. The 
resulting cuttings were removed from the plots. 
Individual T. australe plants were tagged, numbered 
and followed for 2 years (Table 1). 

Although not measured quantitatively at 
Perpendicular Point and Look at Me Now Headland, 
observations indicated that the measurement of bare 
ground may be useful in discussing trends in the data. 
The cover of grasses/herbs and bare ground were 
measured at each census (<5 replicates) in classes (1=1- 

10%, 2=11-20% ,10=91-100%) within randomly 

allocated quadrats ( 1 m 2 ), located within each treatment 



plot. Rock cover was negligible. 

Analysis of the density of T. australe plants in 
each plot (number/100 m 2 ) was made using fully 
factorial ANOVAs and Tukey tests for post hoc 
comparisons. The effects of the treatments (burnt, cut, 
control) were examined on each day of sampling. 

Analyses of the cover classes of bare ground 
were made using a two-way fully factorial ANOVA. 
The effects of treatment (burnt, cut, control) and 
sampling date (pre-treatment, 1 and 2 years post- 
treatment) were examined. 



RESULTS 

Headlands (high density plants) 

Site, but not treatment, had a significant effect 
on the proportion of plants surviving 6 and 16 months 
after the start of the study (Fig. 3). At both times 
survival was higher at Perpendicular Point than at Look 
at Me Now Headland (respectively ^=10, df=l, 
p<0.005; ^=8.5 df=l, p<0.025). By the end of the 
study between 80% and 100% of the original plants 
had suffered mortality. 

Six months after the application of treatments 
there was no significant difference in the density of T. 
australe (0.25 nr 2 ) with respect to treatment and site 
(p>0.05; Fig. 4). Sixteen months after treatment, 
however, there was a significant effect of site (F=4.72, 
df = 1,84, p<0.05). Look at Me Now Headland had a 
higher density of plants than Perpendicular Point. 

At Perpendicular Point there was no significant 
difference in the vigour of T. australe (number of 
stems/plant) with respect to treatment either prior to 
treatment, or 6 months and 16 months after treatment 
(p>0.05; Fig. 5). There appeared to be a general 
increase in plant vigour over this period. 

Within 6 months of applying treatments at 
Perpendicular Point and Look at Me Now Headland, 
flowering and fruiting of original plants and new 
recruits of T. australe were recorded in summer (11 
February 1996). 



60 



Proc. Linn. Soc. N.S.W., 125, 2004 



J.S.COHN 




Date (month and year) 



Figure 3. Proportional survival of T. australe (mean, se) at Perpendicular 
Point (♦) and Look at Me Now Headland (■) following treatments (pooled; 
burnt, cut, control) applied in July 1995. 



a/ Perpendicular Point 
3 -| Treatments applied 



a 

CO 



£ 




b/ Look at Me Now Headland 




Hinterland (low density 
plants) 

The density of T. 
australe plants at Old Bar 
Park was significantly 
affected by treatment 2 years 
after application. The slashed 
areas had a higher density 
than either the burnt or the 
control which were not 
significantly different from 
one another (F=15.5, df=2,3, 
p<0.05; Fig. 6). Pre-treatment 
and 1 year after treatment, 
there was no significant 
difference in the density of T. 
australe between treatments 
(p>0.05). 

There was a 
significant interactive effect 
of treatment and time of 
sampling on the cover of bare 
ground at Old Bar Park 
(F=3.12, df=4, 27, p<0.05; 
Fig. 7). Whilst there was no 
significant difference 
between the plots prior to the 
imposition of treatments, 1 
and 2 years (no significant 
difference) after burning, the 
bare ground was significantly 
higher than in the slashed or 
the control at any time (except 
burn at 2 years = pre-burn and 
cut at 2 years). 

Within 1 year of 
treatment application, 
flowering and fruiting of new 
recruits of T. australe were 
recorded in summer (2 Dec 
1998). 



DISCUSSION 



Date (month and year) 



Figure 4. The density (mean, se) of T. australe plants (0.25 m 2 ) before 
and after treatments at Perpendicular Point and Look at Me Now 
Headland. Treatments (burnt ♦, cut ■, control ▲) were applied in July 
1995. 



Headlands (high density 
plants) 

Burning or cutting T. 
australe plants in winter, did 
not significantly affect their 
survival. Similarly, Leigh and 
Briggs (1989) found the 
survival of a population of T. australe near Canberra, 
was unaffected by a trial burn in autumn. Indeed, the 



Proc. Linn. Soc. N.S.W., 125, 2004 



61 



EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 



a 



10 
8 - 
6 - 
4 

2 H 




Treatments applied 




ON 



ON 



OS 
I 



in 

On 
i 

> 

O 



NO 
On 



NO 
On 



NO 
ON 



NO 
ON 



NO 

ON 

I 

00 



NO 

On 

i 

> 
O 



Date (month and year) 

Figure 5. The size of T. australe plants before and after treatments at 
Perpendicular Point. Size was measured as the number of stems per 
plant (mean, se). Treatments (burnt ♦, cut ■, control A) were applied 
in July 1995. 



b 

o 
o 



o 



15 -| 
10 - 

5 





Treatments 
applied 




1996 1997 1998 

Date (December in year) 

Figure 6. The density (100 m 2 ) of T. australe plants (mean, s.e.) in 
the hinterland at Old Bar Park before and after treatments were 
applied (burnt ♦, slashed ■, control A). Treatments were applied 
in December 1996. 



&3 ^ 

O % 

& J2 

jo o 

U-t s® 

O o^ 

> 

o 
U 



10 - 
8 
6 
4 - 

2 - 




Treatments 
applied 




1996 



1997 



1998 



Date (December in year) 



Figure 7. Cover classes (mean, se) of bare ground before and after 
treatments (burnt ♦, slashed ■, control A), in the hinterland at Old 
Bar Park. Treatments were applied in December 1996. Cover 
classes:l=0-10% 10=11-100%. 



existence of buds in the 
immediate vicinity of the soil 
surface (Mclntyre et al. 1995) 
allows the species to resprout 
after disturbance. In subalpine 
and tableland climates, it is the 
habit of T. australe to die back to 
the rootstock during winter and 
resprout in spring (Cooper 1986; 
Archer 1987; Gross et al. 1995; 
Conn 1999). This is not the case 
in coastal areas, where the species 
persists all year round (Cohn 
1999). 

Whilst a study on the 
southern tablelands of NSW 
(Leigh and Briggs 1989), 
describes T. australe as an annual 
or a biennial, this study suggests 
that the species may live longer 
on the coast. After 6 and 16 
months, respectively, 

approximately 30% and 17% of 
plants were still alive. Since it is 
likely that these plants originated 
at least 9 months previously in 
spring, their ages were more than 
likely 15 months and 25 months, 
respectively. Certainly, Prober 
and Thiele (1998) believe it 
possible that T. australe lives 
longer in less severe climates. 

Although there was no 
significant effect of treatment on 
the density of T. australe, there 
was a higher density at the more 
exposed Look at Me Now 
Headland than at Perpendicular 
Point 2 years after treatments. 
This agrees with Cooper's (1986) 
hypothesis that competition from 
Kangaroo Grass (T. australis) on 
exposed headlands is reduced by 
salt laden winds. It is also possible 
that the experimental burn, which 
was hotter than would be 
experienced naturally, even in 
extreme conditions (R. Bradstock 
pers. coram.), could have led to 
some mortality of T. australe 
seeds near the soil surface, thus 
reducing the effectiveness of this 
treatment. The small size of the 
plots may also have reduced 



62 



Proc. Linn. Soc. N.S.W., 125, 2004 



J.S.COHN 



treatment effectiveness. Finally, more time may have 
been required for the T. australe populations to respond 
to a reduction in competition brought about by the 
experimental treatments. 

Thesium australe is able to grow and reproduce 
very quickly following disturbance in winter. In 
December, 6 months after burning or cutting, there 
was no significant difference in the vigour of plants in 
the treated plots and the control. At the same time 
resprouting plants and new recruits were flowering and 
beginning to fruit. Indeed, flowering and fruiting of T. 
australe at both Perpendicular Point and Look at Me 
Now Headland occurred throughout the year (Conn 
1999). By contrast, flowering and fruiting has been 
found to be seasonal at inland locations, occurring from 
spring to autumn (Stanley and Ross 1983; Briggs and 
Leigh 1985; Gross et al. 1995; Conn 1999). 

Hinterland (low density plants) 

At the more protected Old Bar Park, where T. 
australe was mostly absent from the plots prior to 
treatment, summer slashing rather than burning led to 
significant seedling recruitment of T. australe, 2 years 
after treatment (Fig. 6). Although it is generally 
recognised that burning provides the bare ground for 
seedling establishment that slashing does not (Lunt 
1990a), other factors seemed to be at play in this study. 

The comparable cover of bare ground in all 
treatments at the time of the high numbers of T. australe 
in the slashed plots (Fig. 7), indicates that a reduction 
in grass height may have been responsible. In Victoria, 
Lunt (1990b) believed that selective grazing of 
tussocks to a height of 5 cm by rabbits and kangaroos 
may have contributed to the maintenance of species 
richness by reducing competition from perennial 
grasses. Thus, it is probable that a reduction in the 
height of the dominant species Themeda australis 
rather than an increase in bare ground, led to significant 
recruitment of T. australe seedlings in the slashed 
treatment. 

Given that the post-fire conditions reduced 
competition from Themeda australis, it is curious that 
there was not a significant effect on T. australe 
numbers. In the same summer following burning, 
seedlings of T. australe were observed in these plots 
(S. Long pers. comm.). Their low numbers throughout 
the study, however, may have resulted from the more 
exposed conditions, reflected by higher cover of bare 
ground, experienced during the first and second 
summers (Fig. 7). In addition, T. australe 's 
hemiparasitic dependence on other herbs and grasses, 
(Scarlett et al. 1994), may have made it difficult for T. 
australe to survive, given that its hosts were also 
recovering from the effects of the fire. Indeed summer 



'dying back' of T. australe in times of water stress has 
previously been recorded by Leigh and Briggs (1989). 
Whilst older T. australe plants may have the resources 
to recover, seedlings, such as those observed in this 
experiment soon after the burning, may not have had 
that capability. 

Management implications 

The results from this study, coupled with the 
long-term persistence of T. australe on exposed 
headlands in the absence of active management 
(Cooper 1986; Griffith 1992), indicate that there is 
apparently no need for a change in this regime, except 
where shrub recruitment (native or exotic), may be 
competing with the survival of T. australe. 

By contrast, in the more protected hinterlands, 
where T. australe also occurs, active management may 
be required to reduce competition from Kangaroo 
Grass (T. australis). Although, results from this study 
indicate that early summer slashing of a grassland (5 
cm height) resulted in recruitment of T. australe plants, 
further research is required to determine if this is the 
most appropriate time of the year and method. Of 
particular concern is that disturbance of the summer 
growing Kangaroo Grass (T. australis) at this crucial 
time could result in the introduction of weed species 
(Griffith 1992). There is also a need to determine if 
burning outside summer, especially in autumn or early 
spring, would yield different results. This is important 
in the light of other work, which indicates that fire 
rather than mowing in grasslands is preferable to 
maintain species richness (Kirkpatrick 1986; Lunt 
1991; James 1994). 

If shrub encroachment becomes a threat to the 
survival of T. australe, studies have recommended 
various fire intervals of between 2 and 10 years to 
reduce dominance of native shrubs (Groves 1974; Lunt 
1998b) or a regime of frequent fire and mechanical 
disturbance to reduce exotic shrub frequency (e.g. 
Chrysanthemoides monolifera; Kirkpatrick 1986). 
Although this study did not examine an appropriate 
disturbance interval for T. australe, its quick growth 
and reproductive development and its continued 
presence at Hat Head (S. Griffith pers. comm.), which 
has burnt every 2 to 4 years for the past 15 years (NSW 
NPWS Records), indicates it can apparently cope with 
a relatively frequent disturbance regime. Studies in 
Victoria (Scarlett and Parsons 1982, 1993) suggest that 
the absence of T. australe and other late-flowering 
species along railway lines has resulted from annual, 
late-season burning. Further research is required in 
coastal areas to determine an appropriate disturbance 
interval for the long-term conservation of T. australe. 



Proc. Linn. Soc. N.S.W., 125, 2004 



63 



EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 



ACKNOWLEDGEMENTS 

Thanks to Steve Griffith, Shirley Cohn and Steve 
Clemesha for their assistance in the field. National Parks 
and Wildlife Service staff from Port Macquarie District office 
provided me with encouragement, information and 
equipment. Thanks to Old Bar Bushfire Brigade, who carried 
out the burn at Old Bar Park. Financial assistance was 
provided by Environment Australia and New South Wales 
National Parks and Wildlife Service. Thanks to Andrew 
Denham and Mark Tozer who kindly provided comments 
on the manuscript. 



REFERENCES 

Archer, W.R. (1984). Thesium australe R. Brown 

(Santalaceae)-field notes and observations. The 
Victorian Naturalist 101, 81-85. 

Archer, W.R. (1987). Additional field notes and 

observations of Thesium australe R. Brown 
(Santalaceae). The Victorian Naturalist 104, 46- 
49. 

Bennett, LT. (1994). The expansion of Leptospermum 
laevigatum on the Yanakie Isthmus, Wilson's 
Promontory, under changes in the burning and 
grazing regimes. Australian Journal of Botany 42, 
555-564. 

Briggs, J.D. and Leigh, J.H. (1985). Delineation of 

Important Habitats of Rare and Threatened Plant 
Species in the Australian Capital Territory. 
Unpublished report, National Estate Grants 
Project, Canberra. 

Briggs, J.D. and Leigh, J.H. (1996). 'Rare or Threatened 
Australian Plants'. (CSIRO Publishing: Canberra). 

Cohn, J.S. (1999). Effects of Fire, Mowing and Vertebrate 
Grazing on the Ecology of Thesium australe. 
Endangered Species Project No. 196. Unpublished 
Report, Environment Australia, Canberra. 

Cooper, M.G (1986). A Pilot Survey of 6 Rare Plants in 
NSW. Unpublished Report, NSW NPWS, 
Sydney. 

Crawley, M.J. (1993). 'Methods in Ecology: GLIM for 
Ecologists'. (Blackwell Scientific Publications: 
Melbourne). 

Dodkin, M.J. and Gilmore, A.M. (1985). Species and 

ecosystems at risk: a preliminary review. In 'Bitou 
Bush and Boneseed' (Eds A. Love and R. 
Dyason) pp.33-52. NSW National Parks and 
Wildlife Service and NSW Department of 
Agriculture. 

Griffith, S. J. (1987). A Survey of the Distribution of 
Bitou Bush {Chrysanthemoides monilifera ssp. 
rotundata) on the North Coast of NSW between 
Diamond Head and Seal Rocks, with 
Recommendations for Management. Unpublished 
Report, New South Wales Interdepartmental 
Working Group on Bitou Bush. 

Griffith, S. J. (1992). Species Recovery Plan for Thesium 
australe. Unpublished Report, Endangered 



Species Program, Australian National Parks and 
Wildlife Service, Canberra. 

Gross, C.L., Nano, C„ Jones, R., and Harre, C. (1995). 
Thesium australe (Santalaceae). An Examination 
of Population Sizes at Selected Sites on the New 
England Tablelands, NSW-Summer 1994/95. 
Unpublished Report, Environment Australia, 
Canberra. 

Groves, R.H. (1974). Growth of Themeda australis 
grassland in response to firing and mowing. 
CSIRO, Australia, Division of Plant Industry 
Field Station Record 13, 1-7. 

James, T.A. (1994). Observations on the effects of 

mowing on native species in remnant bushland, 
western Sydney. Cunninghamia 3, 515-519. 

Kirkpatrick, J.B. (1986). The viability of bush in cities-ten 
years of change in urban grassy woodland. 
Australian Journal of Botany 34, 691-708. 

Leigh, J.H. and Briggs, J.D. (1989). Research relating to 
the conservation of rare or threatened plant species 
and their habitats in eastern Australia. In 'The 
Conservation of Threatened Species and their 
Habitats'. (Eds M. Hicks and P. Eiser) pp. 192- 
201. (Occasional Paper No. 2: Canberra). 

Lojnant, B. and Worsoe, E. (1980). The genus Thesium 
Toadflax extinct in Denmark. Flora Fauna 86, 
65-71. 

Long, S.A. (1996). Old Bar Park Plan of Management 
(Draft). (Greater Taree City Council, Taree). 

Lunt, I.D. (1990a). The impact of an autumn fire on a 

long-grazed Themeda triandra (Kangaroo Grass) 
grassland, implications for management of 
invaded, remnant vegetation. The Victorian 
Naturalist 107, 45-51. 

Lunt, I.D. (1990b). Species-area curves and growth-form 
spectra for some herb-rich woodlands in western 
Victoria, Australia. Australian Journal of Ecology 
15, 155-161. 

Lunt, I.D. (1991). Management of remnant lowland 
grasslands and grassy woodlands for nature 
conservation: a review. The Victorian Naturalist 
108, 56-66. 

Lunt, I.D. (1997). Effects of long-term vegetation 
management on remnant grassy forests and 
anthropogenic native grasslands in south-eastern 
Australia. Biological Conservation 81, 287-297. 

Lunt, I.D. (1998a). Two hundred years of land use and 

vegetation change in a remnant coastal woodland 
in southern Australia. Australian Journal of 
Botany 46, 629-647. 

Lunt, I.D. (1998b). Allocasuarina (Casuarinaceae) 

invasion of an unburnt coastal woodland at Ocean 
Grove, Victoria: structural changes 1971-1996. 
Australian Journal of Botany 46, 649-656. 

McDougall, KL. (1989). The Re-establishment of 

Themeda triandra (Kangaroo Grass): Implications 
for the Restoration of Grassland. Technical Report 
Series No. 89, Arthur Rylah Institute for 
Environmental Research, Melbourne. 

McDougall, K.L., and Kirpatrick, J.B. (1994). (Eds) 

'Conservation of Lowland Native Grasslands in 



64 



Proc. Linn. Soc. N.S.W., 125, 2004 



J.S.COHN 



South-eastern Australia'. (World Wide Fund for 
Nature: Sydney). 

Mclntyre, S., Lavorel, S., and Tremont, R.M. (1995). 

Plant life-history: their relationship to disturbance 
in herbaceous vegetation. Journal of Ecology 83, 
31-44. 

McMahon, A.R.G., Carr, G.W., Bedggood, S.E., Hill, 
R.J., and Pritchard, A.M. (1994). Prescribed fire 
and control of coast wattle {Acacia sophorae 
(Labill.) R. Br.) invasion in coastal heath south- 
west Victoria. In 'Fire and Biodiversity: The 
Effects and Effectiveness of Fire Management', 
pp. 87-96. Biodiversity Series Paper No. 8, 
Department of Environment, Sport and 
Territories, Melbourne. 

NPWS (1998). 'Zieria prostrata Recovery Plan'. (NPWS: 
Sydney). 

Parbery, N.H. (1947) Black headland soils of the south 

coast. Unusual process of formation. Agricultural 
Gazette of New South Wales 58, 123-125. 

Prober, S.M. and Thiele, K.R. (1995). Conservation of the 
grassy white box woodlands: relative 
contributions of size and disturbance to floristic 
composition and diversity of remnants. Australian 
Journal of Botany 43, 349-366. 

Prober, S.M. and Thiele, K.R. (1998). Ecology and 

Management of the Austral Toad-flax (Thesium 
australe) at 'Open Grounds', Gillingal Station, 
East Gippsland. Unpublished Report, Department 
of Natural Resources and Environment, Victoria. 

Scarlett, N.H., Bramwell M. and Earl, G. (1994). Austral 
Toad-flax, Thesium australe. Action Statement 
No. 56. Unpublished Report, Department of 
Conservation and Natural Resources, Victoria. 



Scarlett, N.H. and Parsons, R.F. (1982). Rare plants of the 
Victorian Plains. In 'Species at Risk: Research in 
Australia'. (Eds R.H.Groves and W.D.L Ride) pp. 
89-105. (Australian Academy of Science: 
Canberra). 

Scarlett, N.H. and Parsons, R.F. (1990). Conservation 

biology of the southern Australian daisy Rutidosis 
leptorrhynchoid.es. In 'Management and 
Conservation of Small Populations'. (Eds T.W. 
Clark and J.H. Seebeck) pp. 195-205. (Chicago 
Zoological Society: Illinois). 

Scarlett, N.H. and Parsons, R.F. (1993). Rare or 
threatened plants in Victoria. In 'Flora of 
Victoria'. Vol. 1. (Eds D.B. Foreman and N.G. 
Walsh) pp. 227-254. (Inkata Press: Melbourne). 

Stanley, T.D. and Ross, E.M. (1983). 'Flora of South- 
Eastern Queensland'. Volume 1. (Queensland 
Department of Primary Industries: Brisbane). 

Stuwe, J. (1986). An Assessment of the Conservation 

Status of Native Grasslands on the Western Plains, 
Victoria and Sites of Botanical Significance. 
Arthur Rylah Institute Report Series No. 48, 
Department of Conservation, Forests and Lands, 
Melbourne. 

Stuwe, J. and Parsons, R.F. (1977). Themeda australis 

grasslands on the basalt plains, Victoria: floristics 
and management effects. Australian Journal of 
Ecology 2, 467-476. 

Tremont, R.M. and Mclntyre, S. (1994). Natural grassy 

vegetation and native forbs in temperate Australia: 
structure, dynamics and life histories. Australian 
Journal of Botany 42, 641-658. 

Underwood, A.J. (1981). Techniques of analysis of 
variance in experimental marine biology and 
ecology. Annual Reviews of Oceanography and 
Marine Biology 19, 513-605. 



Proc. Linn. Soc. N.S.W., 125, 2004 



65 



66 



Trichromothrips veversae sp.n. (Insecta, Thysanoptera), and the 

Botanical Significance of Insects Host-specific to Austral 

Bracken Fern {Pteridium esculentum) 

Laurence Mound 1 and Mas ami Masumoto 2 

'CSIRO Entomology, GPO Box 1700, Canberra 2601 (laurence.mound@csiro.au); 
2 MAFF, Yokohama Plant Protection Station, Shin'yamashita, 1-16-10, Yokohama, 238-0801, Japan. 

Mound, L. and Masumoto, M. (2004). Trichromothrips veversae sp.n. (Insecta, Thysanoptera), and the 
botanical significance of insects host-specific to Austral bracken fern {Pteridium esculentum). 
Proceedings of the Linnean Society of New South Wales, 125, 67-71. 

Austral bracken fern, Pteridium esculentum, differs from its European counterpart in supporting one 
species of both thrips and aphid. The previously undescribed species of thrips, Trichromothrips veversae 
sp.n. (Thripidae), is widespread and locally abundant in southern Australia breeding on the youngest fronds 
of bracken but not on other ferns. It is unique among nearly 30 species of this Old World tropical genus in 
lacking long setae on the pronotum. 

Manuscript received 18 June 2003, accepted for publication 17 September 2003. 

KEYWORDS: aphids, bracken, Pteridium, thrips, Trichromothrips, 



INTRODUCTION 

Common bracken fern is often considered to 
be a single, cosmopolitan species Pteridium aquilinum 
(Dennstaedtiaceae). In retaining this view, the major 
reference work on botanical nomenclature (Mabberley, 
1997) recognised two subspecies, the nominate one 
from the Northern Hemisphere and Africa, and P. 
aquilinum caudatum from the Southern Hemisphere. 
In Australia, in contrast, Brownsey (1989) recognised 
three species of Pteridium: P. aquilinum introduced 
to a small area of South Australia in the Adelaide Hills; 
P. revolutum native to north-eastern Queensland but 
extending widely across New Guinea and South East 
Asia; and P. esculentum native to southern and eastern 
Australia but extending to South East Asia and the 
Pacific. More recently, Thomson (2000) has concluded 
from an extensive study of both structural and 
molecular characters that several of the Pteridium 
varieties distinguished worldwide, including 
esculentum, "might best be treated as species". 

These differences in opinion concerning the 
botanical status of bracken fern are not without 
entomological significance. No species either of aphid 
(Homoptera) or of thrips (Thysanoptera) is known to 
live on bracken in Europe, where this plant is 
widespread and abundant and often an invasive weed. 
In contrast, the aphid species Shinjia orientalis 
(Mordwilko) (= S. pteridifoliae Shinji) has been 
reported widely on Pteridium from northern India and 
Japan to eastern Australia. Moreover, populations of 



bracken in eastern North America support another 
aphid species, Mastopoda pteridis Oestlund, and in 
western North America five aphid species in the genus 
Macrosiphum have been reported from Pteridium (V. 
F. Eastop, 2003 pers. comm.). If Pteridium were truly 
monotypic, comprising one worldwide panmictic 
species, then different populations might be expected 
to support similar, if not identical insect species. The 
description here of a new species of Thripidae that is 
widespread on bracken in Australia would thus appear 
to provide further support for the recognition of distinct 
species within this ubiquitous plant genus. Presumably 
these insects are reflecting diversity within the genus 
Pteridium that botanists have been reluctant to 
acknowledge. 

The existence of this thrips species had been 
suspected for many years. In 1967, the wife of the 
eminent Australian insect ecologist H.G. Andrewarfha, 
Hattie Vevers-Steele after whom the new species 
described below is named, drew the attention of one 
of us (LAM) to some specimens of a thrips species 
taken from bracken near Adelaide during her studies 
on Australian Thysanoptera (see Mound, 1996). The 
specimens were in poor condition, and efforts at that 
time to locate the species in the field were not 
successful. However, during the past 10 years this 
thrips has been found to be widespread across southern 
Australia, but breeding only in the curled apices of the 
youngest fronds of bracken. This species was listed 
by Shuter and Westoby (1992) from a population of 
bracken near Sydney as "Anaphothripinae gen. et sp. 



THRIPS AND APHIDS ON AUSTRAL BRACKEN FERN 



indet", but is here recognised as a new species of the 
widespread Old World genus Trichromothrips . 
However, within that genus it exhibits one remarkably 
deviant autapomorphy - the absence of any long setae 
on the pronotum. This thrips has been found only on 
Pteridium esculentum, as defined by Brownsey (1998), 
even when this has been found growing in association 
with other ferns that are superficially similar, such as 
the closely related Hypolepis muelleri 
(Dennstaedtiaceae), or young specimens of the more 
distantly related tree fern Dicksonia antarctica 
(Dicksoniaceae). No thrips have been found on any 
species of Hypolepis, although Scirtothrips frondis 
Hoddle and Mound breeds abundantly on the youngest 
fronds of Dicksonia and has also been taken on a 
species of Cyathea (Hoddle and Mound, 2003). 



Trichromothrips Priesner 

Trichromothrips Priesner, 1930: 9. Type species T. 
bellus Priesner. 

Bhatti (2000) has fully defined and reviewed 
this genus, synonymising the genus Dorcadothrips 
Priesner and providing a key to identify the 27 included 
species. Of these, 24 are from the Old World, between 
Africa and Queensland but mostly from South East 
Asia. The other three species, two from Hawaii and 
one widespread, may also have come originally from 
the Oriental region. The collection data for most of 
the species are probably not reliable indicators of the 
plants on which these thrips breed, but two species (71 
billeni Strassen and T. bilongilineatus Girault) are 
associated with ferns (Mound, 2002b), and in the 
region of Japan around Tokyo and Yokahama, T. alis 
Bhatti or a closely related species is found on a species 
of Polystichum (Dryopteridaceae). Finally, three 
related genera of Thripidae are also associated with 
ferns, Laplothrips Bhatti, Octothrips Moulton and 
Pteridothrips Priesner (Mound, 2002b). 

Members of these four genera are unusual in 
bearing a pair of setae on the dorsal apical margin of 
the first antennal segment. This character state is also 
shared by species in the following genera of Thripidae, 
although none involves fern-living species: Alathrips 
Bhatti, Bregmatothrips Hood, Ceratothripoides 
Bagnall, Craspedothrips Strassen, Diarthrothrips 
Williams, Furcithrips Bhatti, Megalurothrips Bagnall, 
Mycterothrips Trybom, Odontothrips Amyot and 
Serville, Odontothripiella Bagnall, Pezothrips Karny, 
Sorghothrips Priesner, Watanabeothrips Okajima, 
Yoshinothrips Kudo. Moreover, although the two 
species comprising the Oriental genus Bathrips Bhatti 
lack this pair of setae on the dorsal apical margin of 



the first antennal segment, they share many other 
character states with Trichromothrips species, and 
these two genera are possibly closely related. 

Trichromothrips veversae sp.n. 

Holotype $ macroptera, Australian Capital 
Territory, Woods Reserve, from young fronds of 
Pteridium esculentum, 6.xii.2002 (LAM 4244), in 
ANIC, CSIRO Entomology, Canberra. 

Paratypes: 2 males, 17 females, same host, 
date and locality as holotype (Masumoto, Mound and 
Wells); 3 females at same locality but 16.1.1999 (LAM 
3664). 

Specimens excluded from the type series 
were collected widely in southern Australia, 
including Tasmania, Western Australia, New South 
Wales, and the Australian Capital Territory (see 
Distribution below). 

Female macroptera 

Colour, body yellow with orange pterothorax, 
ocelli bright red, antennae brown, abdomen with 
transverse light brown markings, wings shaded; colour 
of cleared and mounted specimens yellow, tergites 
shaded anteromedially and along antecostal line, DC 
and X shaded, mesonotum and metanotum weakly 
shaded; head and antennal segment I pale, segments 
m to VIE almost uniformly dark brown with extreme 
base of segments III to V slightly paler, U paler than 
segment HI; all legs greyish brown; fore wing and scale 
greyish brown, but base of fore wing paler. 

Structure: Head slightly wider than long, not 
prolonged in front of eyes, with a few transverse striae 
posteriorly on vertex (Fig. 1); ocellar setae I absent, 
setae III no longer than length of an ocellus and arising 
between anterior margins of posterior ocelli; three pairs 
of postocular setae, pairs I and U close together behind 
ocelli; ventral surface of head with 5 pairs of setae 
between compound eyes anterior to anterior tentorial 
pits; mouth-cone rounded, maxillary palpi 3- 
segmented; compound eyes without pigmented facets. 
Antenna 8-segmented (Fig. 3); forked sense-cones on 
m and IV exceptionally stout; segment I with 2 dorsal 
apical setae; II with weak microtrichia laterally only, 
III to VI with a few large microtrichia on dorsal and 
ventral surfaces; III with 2 dorsal and 2 ventral setae. 

Pronotum medially with few or no lines of 
sculpture and 4 to 10 discal setae; posterior margin 
with five pairs of setae, none of which is longer than 
the discal setae. Mesonotum with weak transverse lines 
of sculpture, without campaniform sensilla near 
anterior margin, median pair of setae far ahead of 
posterior margin. Metanotum (Fig. 2) medially without 
sculpture and one pair of small setae far from anterior 



68 



Proc. Linn. Soc. N.S.W., 125, 2004 



L. MOUND AND M. MASUMOTO 




Figure 1. Trichromothrips veversae, head and 
pronotum. 

margin, without campaniform sensilla. Prosternal ferna 
not divided; mesothoracic sternopleural suture not 
developed; meso- and metasternum each with well- 
developed spinula. All tarsi 2-segmented. Forewing 
veinal setae short, less than half width of wing in 
length; first vein with about 8 setae near base and 2 
(rarely 3) setae near apex; second vein with about 10 
setae; posterior fringe cilia wavy; forewing scale with 
4 marginal setae. 

Abdominal tergites without posteromarginal 
craspeda or lateral ctenidia; tergites II to VIII without 
sculpture medially, lateral to seta S2 with about 7 
anastomosing transverse lines bearing tuberculate 
microtrichia; tergite VIII without posteromarginal 
comb; tergite IX with paired campaniform sensilla 
posteromedially; tergite X undivided; pleurotergites 



without discal seta, sculpture similar to lateral areas 
of tergites. Sternites without discal setae; sternite II 
with two pairs of posteromarginal setae, sternites HI 
to VII with three pairs, on VII all three pairs arise in 
front of sternal posterior margin. 

Measurements (holotype female in |j.m with 
small paratype female in parentheses): Body length 
1400 (1100). Head, length 90 (85); width 125 (105). 
Pronotum, length 105 (95); width 160 (130); 
posteromarginal setae 15 (12). Forewing, length 750 
(650). Antennal segments 25, 32, 50, 57, 40, 43, 10, 
17 (25, 30, 40, 47, 35, 37, 7, 15). 




1 

Y 1 



\ 



Figure 2. Trichromothrips veversae, mesonotum 
and metanotum. 



Male aptera 

Colour paler than female. Structure similar 
to female except: forked sense-cones on antennal 
segments III and IV small and slender; one of three 
available males lacks ocellar setae II; mesonotum 
transverse with 4 or 5 setae near lateral margins; 
pleurotergal sutures weakly developed; tergite IX 



Figure 3. Trichromothrips veversae, antenna. 



«^n; 




^N 



Proc. Linn. Soc. N.S.W., 125, 2004 



69 



THRIPS AND APHIDS ON AUSTRAL BRACKEN FERN 



posterior margin with horn-like paired drepanae 
extending beyond segment X; sternites III to VIII each 
with about 50 small, irregularly arranged, glandular 
areas, marginal setae arising at margin on all sternites. 
Measurements (paratype male in (im). Body 
length 1000. Head, length 83; width 100. Pronotum, 
length 85; width 130; posteromarginal setae 15. Tergite 
IX drepanae length 60. Antennal segments 25, 30, 37, 
40, 32, 37, 7, 15. 

Larva H 

Colour pale yellow with red eyes, 
progressively developing extensive pale red 
hypodermal pigment in meso- and metathorax and 
anterior abdominal segments, body usually turning 
deep yellow progressively; major dorsal setae parallel- 
sided with bluntly square apices, 3 pairs on head, 6 
pairs on pronotum, 3 pairs on abdominal tergites II - 
VIII, 2 pairs on IX, antennal II with 2 pairs of similar 
but smaller setae; setae on tergite X and abdominal 
sternites with apices acute; sternite IX posterior margin 
with row of about 30 small tooth-like tubercles. 

Systematic relationships 

Currently, this new species cannot be placed 
in any of the 10 species-groups distinguished by Bhatti 
(2000) within Trichromothrips, although it shares with 
the other 27 species the many character states listed 
by that author in his diagnosis of the genus. In contrast 
to those species, it lacks any long pronotal setae, the 
metasternal spinula is well developed not weak, and 
females have unusually stout antennal sense cones. 

In Australia, only one other species of 
Trichromothrips has been collected in good numbers: 
T. bilongilineatus (Girault) from ferns near Gosford 
(Mound, 2002a). Of the other two members of the 
genus listed from Australia, the record of T. xanthius 
(Williams) is based on one female taken in quarantine 
in North America but labelled as coming from 
Australia (Mound, 1996), and T. obscuriceps (Girault) 
is known from a single sample apparently taken on 
Crinum lilies near Brisbane. The genus is probably 
well established in northern Australia, but only a few 
specimens are available, representing two further 
unidentified species, swept from grasses near Darwin. 
All of these species have long pronotal posteroangular 
setae. 

The lack of long pronotal setae gives T. 
veversae the superficial appearance of an Anaphothrips 
species. This is another example of the ineffective 
supra-generic classification within the subfamily 
Thripinae, in which traditional subtribal names such 
as Aptinothripina do not refer to definable groups 
(Mound, 2002c), despite their continued use by various 



authors (eg. Vasiliu-Oromulu et al. 2001). There are 
several unrelated Thripinae genera in which species 
usually have two pairs of long pronotal setae, but in 
which one or more species have these setae no longer 
than the discal setae and are thus "Anaphothripine" in 
appearance, eg. Dichromothrips Priesner, 
Pseudanaphothrips Karny and Thrips Linnaeus. 

The presence or absence of long setae on the 
pronotum was recognised as a poor indicator of 
phylogenetic relationships by Mound and Palmer 
(1981), who proposed a series of informal genus- 
groups within the Thripinae. These authors included 
Scolothrips Hinds, a genus of predatory thrips, in their 
Dorcadothrips genus-group (Mound and Palmer, 
1981). Scolothrips species resemble some 
Trichromothrips species in general appearance, for 
example the pale slender body and bulging compound 
eyes, but they have very long ocellar setae and the 
pronotum bears six pairs of elongate setae. Moreover, 
the dorsal apical margin of the first antennal segment 
does not bear a pair of setae, and the mesosternal 
sternopleural sutures are weakly developed. The 
character state on the first antennal segment discussed 
above suggests that the genus-groups recognised by 
Mound and Palmer (1981) require reappraisal. 

Distribution and host records 

T. veversae has been found to be locally 
abundant in many parts of southern Australia, 
including Western Australia near Albany, Tasmania 
near Hobart, and various sites in South Australia 
(Adelaide Hills; Cox's Scrub south of Adelaide; and 
Kangaroo Island). It is abundant in the mountains of 
the ACT, and is widely distributed in the eastern forests 
of New South Wales from near Eden to the Blue 
Mountains. It possibly occurs even further north, but 
a sample taken from Pteridium at Beerwah, north of 
Brisbane, yielded only Scirtothrips dobroskyi Moulton 
(Hoddle and Mound, 2003). In a survey of the insects 
associated with bracken in New Guinea, Kirk (1977) 
does not mention thrips, but since thrips on ferns are 
associated only with very young fronds, or even with 
croziers that are not yet fully expanded, these minute 
insects are often difficult to detect. Similarly, the list 
given by Balick et al. (1978) of insects taken from 
ferns worldwide is based on a survey of published 
records, derived mainly from general collecting, and 
some of the thrips species listed are fungus-feeders, 
not fern-feeders. Mound (2002b) emphasised that 
several published records of thrips on ferns are based 
on single samples or even single specimens, and thus 
cannot be relied on to indicate a host relationship. 

In Japan, the common species of bracken fern 
is considered also to represent Pteridiium esculentum 



70 



Proc. Linn. Soc. N.S.W., 125, 2004 



L. MOUND AND M. MASUMOTO 



and, as indicated above, the aphid species Shinjia 
orientalis has been recorded from this plant in Japan 
as well as Australia. However, searches for thrips on 
substantial populations of bracken in Japan, 
particularly near Narita City, have failed to discover 
Trichromothrips veversae. 

At Crafers in the Adelaide Hills, South 
Australia, a substantial population of adults and larvae 
of Thrips imaginis Bagnall was found on bracken 
fronds in an open field during December 2002, together 
with a few larvae of Trichromothrips veversae. 
However, this seems to be a rare host association for 
the highly polyphagous Australian Plague Thrips. 

At several sites, near Adelaide and on 
Kangaroo Island, larvae of T. veversae were found 
bearing up to 12 larval Eucharitidae (Hymenoptera). 
This is presumably a phoretic association, but no 
observations were made on associated ants, the 
probable host of these small wasps. 



ACKNOWLEDGEMENTS 

The authors are grateful to the Chief, CSIRO 
Entomology, for providing research facilities at Canberra, 
to Victor Eastop of the Natural History Museum, London, 
for information on aphids associated with ferns, and to Alice 
Wells for field assistance and comments on the manuscript. 
John Thomson of the Royal Botanic Gardens Sydney kindly 
drew our attention to several references. Mr. Kenji Morita, 
Head of Yokohama Plant Protection Station, and Mr. Tetsuo 
Imamura, Chief of Identification Section, Yokohama Plant 
Protection Station, kindly facilitated a study visit to Canberra 
by M. Masumoto, and Mr. Shigeo Aochi in Narita City, and 
Mr. Yuji Yoshida in Chiba City provided help in searching 
for thrips on Pteridium in Japan. 



REFERENCES 



Kirk, A. A. (1977). The insect fauna of the weed Pteridium 
aquilinum (L.) Kuhn (Polypodiaceae) in Papua 
New Guinea: A potential source of biological 
control agents. Journal of the Australian 
Entomological Society 16, 403-409 

Mabberley, D.J. (1997). The Plant Book. Second edition. 
Cambridge University Press, Cambridge. 

Mound, L.A. (1996). Thysanoptera, pp 249-336, 397-414 
(Index), in Wells, A., Zoological Catalogue of 
Australia. Volume 26. Psocoptera, Phthiraptera, 
Thysanoptera. Melbourne. CSIRO Australia. 

Mound, L.A. (2002a). Thrips and their host plants: new 
Australian records (Thysanoptera: Terebrantia). 
Australian Entomologist 29, 49-60. 

Mound, L.A. (2002b). Octothrips lygodii sp.n. 

(Thysanoptera, Thripidae) damaging weedy 
Lygodium ferns in south-eastern Asia, with 
notes on other Thripidae reported from ferns. 
Australian Journal of Entomology 41, 216-220. 

Mound, L.A. (2002c). The Thrips and Frankliniella 

genus-groups: the phylogenetic significance of 
ctenidia, pp. 379-386 in Mound L.A. and 
Marullo, R. [eds] Thrips and Tospoviruses. 
Proceedings of the 7 th International Symposium 
on Thysanoptera. Australian National Insect 
Collection, Canberra. 

Mound, L.A. and Palmer, J.M. (1981). Phylogenetic 

relationships between some genera of Thripidae 
(Thysanoptera). Entomologica Scandinavica 15, 
153-17. 

Priesner, H. (1930). Contribution towards a knowledge of 
the Thysanoptera of Egypt, III. Bulletin de la 
Societe Royal Entomologique d'Egypte 14, 6- 
15. 

Thomson, J. A. (2000). Morphological and genomic 
diversity in the genus Pteridium 
(Dennstaedtiaceae). Annals of Botany 85, 77-99. 

Vasiliu-Oromulu, L. zur Strassen, R. and Larsson, H. 

(2001). The systematic revision of Thysanoptera 
species from the Swedish fauna and their 
geographical distribution. Entomologia romana 
6, 93-101. 



Balick, M., Furth, D.G. and Cooper-Driver, G. (1978). 

Biochemical and evolutionary aspects of 

arthropod predation on ferns. Oecologia 35, 55- 

89. 
Bhatti, J.S. (2000). Revision of Trichromothrips and 

related genera (Terebrantia: Thripidae). Oriental 

Insects 34, 1-65. 
Brownsey, P.J. (1989). The taxonomy of Bracken 

{Pteridium: Denstaedtiaceae) in Australia. 

Australian Systematic Botany 2, 113-128. 
Brownsey, P.J. (1998). Denstaedtiaceae, pp. 214-228 in 

Flora of Australia volume 48. Ferns, 

Gymnosperms and Allied Groups. CSIRO, 

Melbourne. 
Hoddle, M.S. and Mound, L.A. 2003. The genus 

Scirtothrips in Australia (Insecta, Thysanoptera, 

Thripidae). Zootaxa 268, 1-40. 



Proc. Linn. Soc. N.S.W., 125, 2004 



71 



72 



Cyst Shell Morphology of the Fairy Shrimps (Crustacea: 

Anostraca) of Australia 

Brian V. Timms 1 , William D. Shepard 2 and Richard. E. Hill 3 

1 School of Environmental and Life Sciences, University of Newcastle, Callaghan NSW 2308. 

2 Department of Biology, California State University, Sacramento California 95819, USA. 

3 3900 Central Avenue, Fair Oaks California 95628, USA. 

Timms, B.V., Shepard, W.D. and Hill, R.E. (2004). Cyst shell morphology of the fairy shrimps (Crustacea: 
Anostraca) of Australia. Proceedings of the Linnean Society of New South Wales 125, 73-95. 

Cyst shell morphology is described for 31 species and 4 genera of endemic Australian fairy shrimp. All 
four genera have distinctive cyst shell structure. 

Manuscript received 3 May 2003, accepted for publication 23 July 2003. 

KEYWORDS: Branchinella, branchiopodid, Parartemia, Streptocephalus. 



INTRODUCTION 

Australia has a relatively simple fauna of 
anostracans, with about 50 species in five genera. The 
list is currently growing because of recent discoveries, 
but presently stands at 29 described and one 
undescribed species of Branchinella 
(Thamnocephalidae), 8 described and 7 undescribed 
species of Parartemia (Parartemidae), two species of 
Artemia (Artemidae), and two described and perhaps 
many undescribed species of Streptocephalus 
(Streptocephalidae) and a species of an undescribed 
branchipodid genus (Timms, in press). All species are 
endemic, except for the two species of Artemia, one 
of which is certainly introduced and the other possibly 
introduced (see McMasters et al., in press; Timms, in 
press). 

Little is known on the biology of Australian 
species (Timms, in press) and even less on detailed 
morphology and of adults and of cysts. Some general 
observations on cyst morphology of some species have 
been made by Herbert and Timms (2000), Timms 
(2002) and Timms and Geddes (2003). 

Cyst shell morphology is much better known 
in fairy shrimp that occur in Europe, Africa and North 
America, and even the species occurring on some 
oceanic islands have been described. Political units 
from which cysts of fairy shrimp have been described 
include: France (Thiery and Gasc 1991); Italy (Mura 
1986, 1991b); California (Hill and Shepard 1997, 
Shepard and Hill 2001); Africa (Brendonck and 
Coomans 1994a, b; Brendonck and Riddoch 1997); 
and the Galapagos Islands of Ecuador (Brendonck et 
al. 1990). Some cyst studies have centered on certain 



families, genera, subgenera or species groups: 
streptocephalids and thamnocephalids (Mura 1992, 
DeWalsche et al. 1991); streptocephalids (Hamer et 
al. 1994a, b); Branchinecta (Mura 1991a), 
Chirocephalus (Mura 2001, Mura et al. 2002), 
Linderiella (Thiery and Chanpeau 1988); 
Streptocephalus (Brendonck et al. 1992; Brendonck 
and Coomans 1994a, b; Streptocephalus 
(Parastreptocephalus) (Brendonck et al. 1992); the 
Streptocephalus sealii group (Shepard 1999). Cyst 
descriptions are also now becoming a common part of 
descriptions of new species. Identification keys based 
solely on cyst morphology have been produced for 
some branchiopod groups (Brendonck and Coomans 
1994a, b; Shepard and Hill 2001; Thiery and Gasc 
1991). Cyst shell morphology is useful for identifying 
all genera so far examined, and many, but not all, 
species. Besides a perceived lack of differences 
between some related species, intraspecific variation 
associated with various predation levels (Mura et al. 
2002, Dumont et al. 2002) and perhaps incipient 
speciation (Mura et al. 2002) may lead to confusion in 
some species. 

Cyst morphology includes both external and 
internal structure (Hill and Shepard 1997). The 
outermost part of the shell is represented by the cortex 
surface. This surface often has spines, spinules, ridges 
of several different forms, buttons and ropey folds. 
The cortex is generally relatively thin. Below the cortex 
is the thick alveolar layer, which may be divided into 
various distinct sublayers. The alveolar layer is usually 
highly vacuolated with the vacuoles separated by 
curved or straight, solid struts of varying length. The 
innermost layer is the tertiary base which is relatively 
thin, solid and sometimes lamellate throughout. Often 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 



the interior surface has an impressed polygonal pattern. 
An additional cyst shell character is inpocketing of 
the entire shell. Inpocketing is only typical of cyst shells 
which have a fairly uniform alveolar layer. 

The goal of this paper is to describe the 
external and internal structure of the cyst shell in as 
many as possible of the endemic Australian fairy 
shrimp fauna. This is a prerequisite for understanding 
differences between genera and species, as a base for 
understanding any intraspecific differences and for the 
construction of identification keys. Comparative 
information on Anemia cysts is provided by Shepard 
and Hill (2001) and is not repeated here. This is of 
overseas material, but is unlikely to be different from 
Australian cysts, as all Anemia cysts are the same (G. 
Mura, pers. com.; W. Shepard, pers. com.), except for 
A. monica, which does not occur here. 



METHODS 

Specimens of fairy shrimps were collected 
in the field and preserved in 70-80% ethyl alcohol and 
identified by BVT using Geddes 1981 and Timms in 
press. Females with the most mature looking cysts 
(those with a regular pattern all over the surface and 
which retain their sphericity) present in the brood 
pouch(es) were selected for dissection of the cysts. 
Cysts were removed and air-dried on filter paper. Then 
the cysts were mounted on SEM stubs using double- 
stick tape and gold-coated. A Zeiss DSM 940 SEM 
was used for measuring and photographing the cysts. 
Cyst morphology is described as in Hill and Shepard 
(1997), Shepard (1999) and Shepard and Hill (2001). 
Cyst stubbs have been stored in the private collection 
ofREH. 



RESULTS 

Cysts were obtained for 3 1 of the 50 or so 
species of fairy shrimp known to exist in Australia. 
Those species include: Branchinella affinis Linder 
1941, B. arbor ea Geddes 1981, B. australiensis 
(Richters 1876), B. basipina Geddes 1981, B. 
buchananensis Geddes 1981, B. budjiti Timms 2001, 
B. campbelli Timms 2001, B. complexidigitata Timms 
2002, B. dubia (Schwartz 1917), B. frondosa Henry 
1924, B. hattahensis Geddes 1981, B. kadjikadji 
Timms 2002, B. lamellata Timms and Geddes 2003, 
B. longirostris Wolf 1911,5. lyrifera Linder 1941, B. 
nana Timms 2002, B. nichollsi Linder 1941, B. 
occidental is Dakin 1914, B. pinnata Geddes 1981, B. 
proboscida Henry 1924, B. simplex Linder 1941, B. 



wellardi Milner 1929, Parartemia contracta Linder 
1941, P. cylindrifera Linder 1941, P. informis Linder 
1941, P. minuta Geddes 1973, P. zietziana Sayce 1903, 
Streptocephalus queenslandicus Herbert and Timms 
2000, Streptocephalus n. sp. a, Streptocephalus n. sp. 
b and an undescribed branchiopodid anostracan (all 
three from the Paroo, nw NSW-sw Qld - Timms and 
Sanders 2002 and Timms unpublished data. Two 
different cyst types were found in Branchinella 
occidentalis. Full descriptions could not be made for 
Branchinella affinis, B. nana and Parartemia informis 
due to the presence of only immature cysts. Locality 
and collection details for each species are listed in 
Appendix 1. The described Australian fairy shrimp 
species for which we did not have cysts include the 
following: Branchinella apophystata Linder 1941, B. 
compacta Linder 1941, B. denticulata Linder 1941, 
B. halsei Timms 2002, B. insularis Timms and Geddes 
2003, B. latzi Geddes 1981, B. tyleri Timms and 
Geddes 2003, Parartemia extracta Linder 1941, P. 
longicaudata Linder 1941, P. serventyi Linder 1941 
and Streptocephalus arched Sars 1896. 

The cyst characteristics of Australian 
anostracans are as follows. 

Family Parartemidae 

Parartemia contracta Linder 

Cyst shape spherical (Fig. 1). Cyst size 250- 
275 um (mean = 263.3; n = 10). External surface with 
some inpocketing, adjacent inpockets separated by 
indistinct ridges; individual cysts with 2-6 inpockets; 
surface smooth. Alveolar layer (Fig. 2) uniform; short 
struts; vesicles small rounded and subequal. Tertiary 
layer thin. 

Parartemia cylindrifera Linder 

Cyst shape spherical (Fig. 3). Cyst size 204- 
225 um (mean = 214.6; n = 10). External surface with 
some inpocketing; adjacent inpockets separated by 
ridges; individual cysts with several inpockets each; 
surface smooth. Alveolar layer uniform, mostly solid 
(Fig. 4). Tertiary layer thin. 

Parartemia informis Linder 

Immature cyst shape spherical. Cyst size 162- 
176 um (mean = 167.5; n = 10). 

Parartemia minuta Geddes 

Cyst shape spherical (Fig. 5). Cyst size 180- 
232 um (mean = 208.0; n = 30). External surface with 
inpocketing; adjacent inpockets separated by ridges; 
individual cysts with several inpockets each; surface 
smooth. Alveolar layer uniform and solid (Fig. 6). 
Tertiary layer thin. 



74 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 



Parartemia zietziana Sayce 

Cyst shape spherical to hemispherical (Figs. 
7-8). Cyst size 180-208 [mi (mean = 192.7; n = 11). 
External surface with inpocketing; inpocketing usually 
at one end; adjacent inpockets separated by ridges; 
ridges usually with sloping walls and rounded tops; 
surface smooth. Alveolar layer uniform and solid (Fig. 
9). Tertiary layer thin. 

Family Branchiopodidae 

undescribed branchiopod genus and species 

Cyst shape very irregularly spherical (Fig. 
10), with projecting flanges that interlock with those 
of other cysts binding the cyst mass together (Fig. 11). 
Cyst diameters 208-250 urn (mean = 229.7; n = 20). 
External surface with ridges defining irregular 
polygons; intersecting ridges often extending as 
flanges; entire surface covered with shallow, rounded 
to polygonal depressions; polygon size variable and 
larger on flanges and ridges (Fig. 12). Cortex relatively 
thick (» 2 \ym); one layer of short columns, columns 
parallel to each other and perpendicular to the surface. 
Alveolar layer (Fig. 13) thin; with small and uniform 
vesicles. Tertiary base thin. 

Family Streptocephalidae 

Streptocephalus queenslandicus Herbert and Timms 
Cyst shape tetrahedral (Fig. 14-15); cyst size 
169-208 \im (mean = 195; n = 20). External surface 
with ridges dividing surface into 4 triangular faces; 
ridges each with 2 longitudinal grooves in middle half 
producing 3 short ridge crests; intersecting ridges flare 
outward; triangular faces gently convex with 1 or 2 
small bumps midface; surface granular. Alveolar layer 
(Fig. 16) with 2 sublayers; inner sublayer with short 
straight struts and medium-sized vesicles and hollows; 
outer sublayer uniform and solid; vesicular layer 
extends outward into ridge intersections. Tertiary layer 
thin. 



Streptocephalus n. sp. b (locality: Box Hole, on road 
3 km south of homestead, Currawinya National Park, 
SW QLD) 

Cyst shape tetrahedral (Fig. 20); cyst size 240- 
268 \xm (mean = 249.3; n = 20). External surface with 
ridges dividing surface into 4 triangular faces; ridges 
each with 2 longitudinal grooves in middle producing 
3 short ridge crests (Fig. 21); intersecting ridges 
producing rounded corners; triangular faces depressed, 
flattened, with small bumps midface; surface mildly 
bumpy. Alveolar layer (Fig. 22) vesicular; inner 
vesicles smaller; vesicles under ridges larger. Tertiary 
layer thin. 

Family Thamnocephalidae 

Branchinella affinis Linder 

Cyst shape spherical (Fig. 23). Cyst diameter 
95-134 |j,m (mean = 113.2; n = 20). External surface 
with steep-walled, arcuate ridges defining pinched 
polygons. 

Branchinella arborea Geddes 

Cyst shape spherical (Fig. 24). Cyst size 183- 
201 (Am (mean = 191.6; n = 10). External surface with 
ridges defining flat-bottomed polygons; ridges straight- 
sided, surface covered with small punctae (Fig. 25). 
Alveolar layer (Fig. 26) with 2 sublayers; inner 
sublayer with small subequal vesicles; outer sublayer 
with long struts and large hollows. Tertiary base thin. 

Branchinella australiensis (Richters) 

Cyst shape spherical (Fig. 27). Cyst size 197- 
222 um (mean = 213.5; n = 10). External surface with 
ridges defining polygons; ridges straight- walled, top 
pinched into a bead running along ridge, bead with 
pores (Fig. 28); polygons with flat bottoms. Alveolar 
layer (Fig. 29) with variably-sized vesicles; inner 
vesicles usually smaller than outer ones. Tertiary base 
thin. 



Streptocephalus n. sp. a (locality: Pine Tree Pool, 
Budgerie Paddock, Bloodwood Station, 130 km NW 
of Bourke, NSW) 

Cyst shape tetrahedral (Fig. 17); cyst size 254- 
300 [im (mean = 283.0; n = 20). External surface with 
ridges dividing surface into 4 triangular faces; ridges 
with nearly vertical sides, top rounded (Fig. 18), middle 
half of ridge along each face with a series of large 
punctae on each side, intersecting ridges flare outward; 
triangular faces flat to convex; surface bumpy. Cortex 
thick (2-3 (Am). Alveolar layer (Fig. 19) with 2 
sublayers; inner sublayer with thin short struts defining 
variably-sized, interconnecting vesicles; outer sublayer 
uniform and solid. Tertiary layer thin. 



Branchinella basipina Geddes 

Cyst shape spherical (Fig. 30). Cyst size 225- 
275 um (mean = 253.5; n = 10). External surface with 
ridges defining pinched polygons; ridges with sloping 
sides, tops evenly rounded; polygons with concave 
bottoms; surface rugose. Alveolar layer (Fig. 31) with 
small, subequal vesicles. Tertiary base thin. 

Branchinella buchananensis Geddes 

Cyst shape spherical (Fig. 32). Cyst size 204- 
232 fun (mean = 217.9; n = 10). External surface with 
ridges defining pinched polygons; ridges with sloping 
sides; polygons with concave bottoms; surface rugose. 
Alveolar layer (Fig. 33) with 2 sublayers; inner 



Proc. Linn. Soc. N.S.W., 125, 2004 



75 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 



sublayer with small subequal vesicles; outer sublayer 
with long branching struts and large unequal vesicles. 
Tertiary base thin. 

Branchinella budjiti Timms 

Cyst shape spherical (Fig. 34). Cyst size 141- 
155 um (mean = 144.7; n = 23). External surface with 
sinuous ridges defining tightly pinched polygons; 
ridges steep-sided with rounded tops; polygons with 
sinuous valley-like bottoms; surface slightly bumpy. 
Alveolar layer (Fig. 35) with 2 sublayers; inner layer 
with small vesicles; outer sublayer with long parallel 
struts and large hollows. Tertiary layer thin. 

Branchinella campbelli Timms 

Cyst shape spherical (Fig. 36). Cyst size 162- 
194 um (mean = 172.3; n = 20). External surface with 
broad ridges defining pinched polygons; ridges with 
sides sloped to vertical, ridge tops very broad; surface 
scaley. Alveolar layer (Fig. 37) thick, with small to 
moderate sized vesicles. Tertiary layer thin. 

Branchinella complexidigitata Timms 

Cyst shape spherical (Fig. 38). Cyst size 21 1- 
307 um (mean = 251.0; n = 40). External surface with 
ridges defining irregular polygons; ridges narrow, 
steep-sided, midline of ridges with sharp-pointed 
projections (Figs. 39-40); polygons with bottom flat 
to concave; surface smooth to scaley and porous. 
Alveolar layer (Fig. 41) thin; small to medium sized 
circular vesicles. Tertiary base thin. 

Branchinella dubia (Schwartz) 

Cyst shape spherical (Fig. 42). Cyst size 187- 
215 um (mean = 197.1; n = 32). External surface with 
ridges defining polygons; ridges straight, steep-walled 
and narrow; polygons with bottoms flat to slightly 
concave; surface covered with short wrinkles, pores 
abundant (Fig. 43). Alveolar layer (Fig. 44) with 2 
sublayers; inner sublayer with small rounded subequal 
vesicles; outer sublayer with long sometimes branching 
struts and large hollows. Tertiary base thin. 

Branchinella frondosa Henry 

Cyst shape spherical (Fig. 45). Cyst size 185- 
21 1 um (mean = 196. 1 ; n = 10). External surface with 
ridges defining polygons; ridges with sloping sides, 
tops rounded; polygons with bottom flat to slightly 
concave; surface smooth. Alveolar layer (Fig. 46) with 
3 sublayers; inner sublayer with small vesicles; middle 
sublayer with long sometimes branching struts and 
large hollows; outer sublayer with medium-sized 
vesicles. Tertiary base thin. 



Branchinella hattahensis Geddes 

Cyst shape spherical (Fig. 47). Cyst size 254- 
289 um (mean = 268.9; n = 20). External surface with 
ridges defining polygons; ridges with steep sloping 
sides, ridge tops with a sinuous bead and sharp-pointed 
and sometimes recurved projections (projections 
sometimes with film between) (Figs. 48-49); polygons 
with bottoms flat to slightly concave; surface strongly 
rugose or bumpy. Alveolar layer (Fig. 50) with 2 
sublayers; inner sublayer with small- to medium-sized 
rounded vesicles; outer sublayer with narrow curving 
struts and large hollows. Tertiary base thin. 

Branchinella kadjikadji Timms 

Cyst shape spherical (Fig. 51). Cyst size 254 
um (n = 2). External surface with ridges defining 
polygons; ridges steep-sided and sinuous, tops with 
rounded bead; polygons irregularly shaped, bottoms 
flat; surface smooth (Fig. 52). Alveolar layer (Figs. 
53-54) with at least 2 sublayers; outer sublayer with 
medium-sized rounded vesicles; next sublayer inside 
with short struts and large hollows. Tertiary base not 
visible in available cross-sections. 

Branchinella lamellata Timms & Geddes 

Cyst shape spherical (Fig. 55). Cyst size 124- 
180 [Am (mean = 147.6; n = 31). External surface with 
ridges defining polygons; ridges with sloping sides and 
rounded tops; polygons irregular and highly pinched, 
bottoms concave; surface bumpy. Alveolar layer (Fig. 
56) with narrow struts and very large hollows. Tertiary 
base thin. 

Branchinella longirostris Wolf 

Cyst shape spherical (Fig. 57). Cyst size 264- 
300 -um (mean = 276.9; n = 29). External surface with 
ridges defining polygons; ridges very narrow, not 
always connecting to other ridges, ridge junctions 
extended into spines with tips that are bifid, trifid or 
recurved (Fig. 58); polygons with concave bottoms; 
surface bumpy to scaley. Alveolar layer (Fig. 59) 
mostly hollow with very narrow struts. Tertiary base 
thin. 

Branchinella lyrifera Linder 

Cyst shape spherical; inpocketing common 
(Figs. 60). Cyst size 158-183 um (x = 171.5; n = 20). 
External surface with numerous short flat-topped 
columns, columns often connected by ridges; ridges 
of variable widths (Figs. 61-62). Alveolar layer solid 
(Fig. 63). Tertiary base thin. 

Branchinella nana Timms 

Immature cyst shape spherical . Cyst size 144- 
158 um (mean = 152.3; n = 19). 



76 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 



Branchinella nichollsi Linder 

Cyst shape spherical (Fig. 64). Cyst size 187- 
247 um (mean = 202.3; n = 30). External surface with 
ridges defining polygons; ridges narrow, steep-sided, 
with zig-zag patterns, tops with narrow bead (Fig. 65); 
polygons tightly pinched, bottoms flat to concave; 
surface slightly bumpy. Alveolar layer (Fig. 66) mostly 
hollow, with very thin struts. Tertiary base thin. 

Branchinella occidentalis Dakin (two cyst morphs) 

Type I cyst: Cyst shape spherical; inpocketing 
common (Fig. 67). Cyst size 257-328 um (mean = 
283.5; n = 20). External surface with narrow sinuous 
ridges defining small, irregular, pinched polygons; 
ridges steep-sided and short. Alveolar layer (Fig. 68) 
not well developed. Tertiary base not seen. 

Type II cyst: Cyst shape spherical (Fig. 69). 
Cyst size 550-571 um (mean = 565.3; n = 20). External 
surface with ridges defining rounded depressions; 
ridges narrow except where intersecting, sides vertical, 
tops flat; rounded depressions with bottoms slightly 
concave and extremely porous (Fig. 70); surface 
smooth. Alveolar layer (Fig. 71) with 3 sublayers; inner 
sublayer with small equal rounded vesicles; middle 
sublayer with few struts, mostly open hollows, pores 
connecting to outside; outer layer with stringy 
vermiform struts, connecting to inner side of external 
surface. Tertiary layer thin. 

Branchinella pinnata Geddes 

Cyst shape spherical (Fig. 72). Cyst size 173- 
190 um (mean =181.1;n=10). External surface with 
ridges defining pinched polygons; ridges with sides 
nearly vertical, tops rounded; polygons irregular in 
shape, bottoms flat to concave; surface very bumpy. 
Alveolar layer (Fig. 73) with 3. sublayers; inner 
sublayer with small rounded subequal vesicles; middle 
layer with long branching struts and large hollows; 
outer sublayer with medium-sized irregularly shaped 
vessicles. Tertiary layer thin. 

Branchinella probiscida Henry 

Cyst shape spherical (Fig. 74). Cyst size 158- 
187 um (mean = 174.9; n = 19). External surface with 
ridges defining polygons; ridges vermiform with sides 
nearly vertical, tops smooth and rounded; polygons 
irregular in shape, some pinched, bottoms slightly 
concave and with pores; surface generally smooth (Fig. 
75). Alveolar layer (Fig. 76) without distinct sublayers; 
struts usually short; vesicles irregular in shape and 
variable in size. Tertiary layer thin. 

Branchinella simplex Linder 

Cyst shape spherical (Fig. 77). Cyst size 144- 



201 um (mean = 176.4; n = 32). External surface with 
ridges defining polygons; ridges with sides slightly 
sloping to vertical; polygons pinched, bottoms 
concave; surface smooth with irregularly spaced 
punctae. Alveolar layer (Fig. 78) uniform; short 
branching struts and medium-sized hollows. Tertiary 
layer thin. 

Branchinella wellardi Milner 

Cyst shape spherical (Fig. 79). Cyst size 158- 
176 um (mean = 168.4; n = 30). External surface with 
ridges defining polygons; ridges narrow and steep- 
sided, tops sharp except broader at intersections (Fig. 
80); polygons irregular in shape, some pinched, 
bottoms concave to flat; surface punctate to porous. 
Alveolar layer (Fig. 81) uniform, thick; short struts 
with small interconnected vesicles. Tertiary layer thin. 



DISCUSSION 

Australia is currently experiencing a surge of 
interest in its fairy shrimps, as indicated by the large 
percentage of newly described species and species 
awaiting description (Timms, in press). Species 
descriptions omit information on cyst shells, but 
suitable descriptions for many species are provided 
here. Although about 40% of known species are yet to 
be studied (and for most of these, data are unlikely to 
be available in the near future), enough descriptions 
of cyst shell morphology are available for comparison 
between Australian and overseas fauna and among the 
Australian species. 

Cysts of the species examined could be easily 
separated at the genus level, but only with difficulty 
or not at all at the species level. The most distinctive 
are those of Streptocephalus with their overall 
tetrahedral shape and distinct facet-edge ridges and 
convex to concave faces with bumps. This feature 
places them within the sudanicus species group of 
Maeda-Martinez et al. (1995) and subgenus 
Parastreptocephalus of Brendonck et al. (1992); 
Streptocephalus species with mature spherical cysts 
have yet to be found in Australia. 

Cysts of the new branchiopodid genus are 
also distinctive because of flanges from ridges defining 
irregular polygons. Almost all of the species of 
Branchinella have cysts with a polygonal surface 
largely reminiscent of those of overseas members of 
the genus (Belk and Sisson 1992, Mura 1992, 
Brendonck and Riddoch 1997; Sanoamuang et al. 
2003). Cysts of B. lyrifera are discordant for the genus 
because they are covered with flat-topped columns 
with little evidence of polygons, though the columns 



Proc. Linn. Soc. N.S.W., 125, 2004 



77 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 



are connected by uncoordinated ridges. 

Cysts of Parartemia, an Australian endemic 
genus, are described for the first time. They are smooth, 
rather like those of Anemia (Hill and Shepard 1997), 
but with some inpocketing. Given possible undetected 
variation within species, particularly in those subjected 
to predation (Mura et al. 2002), it is unwise to compare 
species within genera. 

None of the Australian cysts examined here 
have extreme anti-predation devices such as 
honeycombing or long spines (Dumont et al. 2003). 
Some species have minor protruding structures which 
may assist against predation. The most developed are 
the bifid/trifid/recurved spines at ridge junctions in B. 
longirostris. Interestingly this species lives in gnammas 
(rock pools) in which many potential predators live 
including flatworms (Pinder et al. 2000). Other species 
of Branchinella with small spines include B. 
complexidigitata in which they are sharp pointed and 
B. hattahensis where the spines are recurved. The 
flanges on the cysts of the undescribed branchipodid 
genus could be antipredation devices, but another 
explanation is that they assist in holding the egg mass 
together (as observed when trying to separate the 
cysts). The value of such a strategy is questionable, 
given the decreased possibly of dispersal of a clumped 
egg mass, but perhaps it helps to deter egg predators. 

The cyst shell structure (especially the cross- 
sectional structure) in this study ranged from relatively 
simple (e. g., in Parartemia) to complex (e. g., in 
Branchinella budjiti and an undescribed branchipodid 
genus). During the examination of Branchinella 
occidentalis the first set of cysts appeared to be not 
completely mature so a second set was examined. This 
second set was of a very different size and form. Thus, 
it appears that either the species has two different cyst 
morphs or there may be a cryptic species in what is 
now known as B. occidentalis. At least other species 
are known to have two cyst types: Eubranchipus 
serratus (Hill and Shepard 1997) and Streptocephalus 
sealii (R. Hill, pers.com.). During the course of this 
study the only cyst structure found that has not been 
seen in any other fairy shrimp before was the external 
flanges and the parallel columns in the cortex of the 
undescribed genus of branchipodid fairy shrimp. 
Therefore, with minor exception, the maximum range 
of structural diversity in cysts may have been 
described. Whether or not this is true will be decided 
by examination of the remaining undescribed cysts of 
other world genera. 

The structure of the cyst shell, especially in 
the alveolar layer, begs a functional explanation. It is 
likely that the structure represents a compromise 
between crush-resistance and rehydration ability. But 
other functions that have been suggested include: 



protection from abrasion and ultraviolet light (Belk 
1969); a float for cysts (Morris and Afzelius 1967); a 
barrier against physio-chemical stresses and an aid to 
gaseous exchange (Tommasini and Sabelli 1989); an 
anti-predation device (Dumont et al. 2003) and, 
dispersal aids or inhibitors (Brendonck et al. 1992). 



ACKNOWLEDGEMENTS 

We thank Bruce Scott for his masterful supervision of 
the scanning electron microscope. 



REFERENCES 

Belk, D.. (1969). Functions of the conchonstracan eggshell. 
Crustaceana 19(1), 106-107. 

Belk, D. and Sissom, S. (1992). New Branchinella 
(Anostraca) from Texas, U.S.A., and the problem 
of antennalike processes. Journal of Crustacean 
Biology 12, 312-316. 

Brendonck, L. and Coomans, A. (1994a). Egg morphology 
in African Streptocephalidae (Crustacea: 
Branchiopoda: Anostraca) Part 1: South of 
Zambezi and Kunene rivers. Archiv 
Hydrobiologia/Supplement 99 (3), 313-334. 

Brendonck, L. and Coomans, A. (1994b). Egg morphology 
in African Streptocephalidae (Crustacea: 
Branchiopoda: Anostraca) Part 2: North of 
Zambezi and Kunene rivers, and Madagascar. 
Archiv Hydrobiologia/Supplement 99 (3), 335- 
356. 

Brendonck, L. and Riddcoh, B. (1997). Anostracans 
(Branchiopoda) of Botswana: Morphology, 
Distribution, Diversity, and Endemicity. Journal 
of Crustacean Biology 17, 111-134. 

Brendonck, L., Hamer, M. and Thiery, A. (1992). Occurrence 
of tetrahedral eggs in the Streptocephalidae Daday 
(Branchiopoda: Anostraca) with descriptions of a 
new subgenus, Parastreptocephalus, and a new 
species, Streptocephalus (Parastreptocephalus) 
zuluensis Brendonck and Hamer. Journal 
Crustacean Biology 12, 282-297. 

Brendonck. L., Thiery, A. and Coomans, A. (1990). 
Taxonomy and biogeography of the Galapagos 
branchiopod fauna (Anostraca, Notostraca, 
Spinicaudata). Journal Crustacean Biology 10, 
676-694. 

Dakin, W.J. (1914). Fauna of Western Australia. II The 
Phylllopoda of Western Australia. Proceedings 
Zoological Society London 1914, 293-305. 

DeWalsche, C. Munuswamy, N. and Dumont, H. J. (1991). 
Structural differences between the cyst walls of 
Streptocephalus dichotomus (Baird), S. torvicornis 
(Waga), and Thamnocephalus platyurus (Packard) 
(Crustacea: Anostraca), and a comparison with 
other genera and species. Hydrobiologia 212, 195- 
202. 



78 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 



Dumont, H.J., Nandini, S. and Sarma, S.S.S. (2003). Cyst 
ornamentation in aquatic invertebrates: a defence 
against egg-predation. Hydrobiologia 486, 161- 
167. 

Geddes, M.C. (1973). Studies on Australian Anostracans 
(Crustacea: Branchiopoda). Ph. D. thesis. Monash 
University. 282 pp. 

Geddes, M.C. (1981). Revision of Australian species of 
Branchinella (Crustacea: Anostraca). Australian 
Journal of Marine and Freshwater Research 32, 
253-295. 

Hamer, M., Brendonck, L. Coomans, A. and Appleton, C. 
C. (1994). A review of African Streptocephalidae 
(Crustacea: Branchiopoda: Anostraca) Part 1: 
South of Zambezi and Kunene rivers. Archiv 
Hydrobiologia/Supplement 99 (3), 235-277. 

Hamer, M., Brendonck, L. Coomans, A. and Appleton, C. 
C. (1994). A review of African Streptocephalidae 
(Crustacea: Branchiopoda: Anostraca) Part 2: 
North of Zambezi and Kunene rivers and 
Madagascar. Archiv Hydrobiologia/Supplement 99 
(3), 279-311. 

Henry, M. (1924). A monograph on the freshwater 
Entomostraca of New South Wales. IV 
Phyllopoda. Proceedings Linnean Society New 
South Wales 49, 120-137. 

Herbert, B. and Timms, B.V. (2000). A new species of 
Streptocephalus (Parastreptocephalus) 
(Crustacea: Anostraca: Streptocephalidae) from 
North Queensland, Australia. Memoirs of the 
Queensland Museum 45(2), 385-390. 

Hill, R.E. and Shepard, W.D. (1997). Observations on the 
identifications of California anostracan cysts. 
Hydrobiologia 359, 113-123. 

Linder, F. (1941). Contributions to the morphology and the 
taxonomy of the Branchiopoda Anostraca. 
Zoologiska Bidrag frdn Uppsala 20, 101-302. 

Maeda-Martinez, A.M., Belk, D., Obregon-Barboza, H and 
Dumont, H.J. (1995). A contribution to the 
systematics of the Streptocephalidae 
(Branchiopodidae: Anostraca). Hydrobiologia 
298, 203-232. 

McMaster, K., Savage, A., Finston, T. Johnson, M.S. and 
Knott, B. (In Press). Has Artemia parthenogenetica 
been introduced into Western Australia through 
human agency? Hydrobiologia 

Milner, D.F. (1929). A description of two new species of 
Anostraca Phyllopoda from Western Australia. 
Journal Royal Society of Western Australia 15, 
25-35. 

Morris, J. E. and Afzelius, B. A. (1967). The structure of 
the shell and outer membranes in encysted Artemia 
salina embryos during cryptobiosis and 
development. Journal Ultrastructure Research 20, 
244-259. 

Mura, G. (1986). SEM morphological survey on the egg 
shell of Italian anostracans (Crustacea, 
Branchiopoda). Hydrobiologia 134, 273-286. 

Mura, G. (1991a). SEM morphology of resting eggs in the 
species of the genus Branchinecta from North 



America. Journal Crustacean Biology 11, 432- 
436. 

Mura, G. (1991b). Additional remarks on cyst 
morphometries in anostracans and its significance. 
Part I: egg size. Crustaceana 61, 241-252. 

Mura, G. (1992). Pattern of egg shell morphology in 
thamnocephalids and streptocephalids of the New 
World (Anostraca). Crustaceana 62, 300-311. 

Mura, G. (2001). Morphological diversity of the resting eggs 
in the anostracan genus Chirocephalus (Crustacea, 
Branchiopoda). Hydrobiologia 450, 173-185. 

Mura, G., Zarattini, P. and Petkowski, S. (2002). 
Morphological divergences and similarities among 
Chirocephalus diaphanous carinatus populations 
(Crustacea, Anostraca) from the Balkan area. 
Journal of Crustacean Biology 22, 162-172. 

Pinder, A.M., Halse, S.A., Shiel, R.J. and McRae, J.M. 
(2000). Granite outcrop pools in south-western 
Australia: foci of diversification and refugia for 
aquatic invertebrates. Journal of the Royal Society 
of Western Australia 83, 149-161. 

Richters, F. (1876). Branchipus australiensis n. sp. Journal 
Museum Godeffroy 12, 43-44. 

Sanoamuang, L., Saenghan, N, and Murugan, G. (2003). 
First record of the family Thamnocephalidae 
(Crustacea: Anostraca) from Southeast Asia and 
description of a new species of Branchinella. 
Hydrobiologia 486, 63-69. 

Sars, G.O. (1896). Description of two new Phyllopoda from 
north Australia. Archiv Mathematik Naturvidensk 
Christiana 28, 4-6. 

Sayce, O.A. (1903). The Phyllopoda of Australia, including 
descriptions of some new genera and species. 
Proceedings Royal Society Victoria 15, 224-261. 

Schwartz, K.Y. (1917). Results of Dr. E Mjobergs Swedish 
scientific expedition to Australia 1910-1913. XV. 
Descriptions of two Australian phyllopods. K. 
Sven. Vetenkapsakad. Handl. 52, 7-8. 

Shepard, W.D. (1999). First record of fairy shrimp in Belize, 
and a comparison of cyst-shell morphology in the 
New World members of the Streptocephalus sealii 
species group (Anostraca: Streptocephalidae). 
Journal of Crustacean Biology 19, 355-360. 

Shepard, W.D. and Hill, R.E. (2001). Anostracan cysts found 
in California salt lakes. Hydrobiologia 466, 149- 
158. 

Thiery, A. and Champeau, A. (1988). Linderiella 
massaliensis, new species (Anostraca: 
Linderiellidae), a fairy shrimp from south-eastern 
France, its ecology and distribution. Journal 
Crustacean Biology 8, 70-78. 

Thiery, A. and Gasc, C. (1991). Resting eggs of Anostraca, 
Notostraca and Spinicaudata (Crustacea, 
Branchiopoda) occurring in France: identification 
and taxonomical value. Hydrobiologia 212, 245- 
259. 

Timms, B.V. (2001). Two new species of fairy shrimp 
(Crustacea: Anostraca: Thamnocephalidae: 
Branchinella) from the Paroo, inland Australia. 
Records Australian Museum 53, 247-254. 



Proc. Linn. Soc. N.S.W., 125, 2004 



79 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 



Timms, B.V. (2002). The fairy shrimp genus Branchinella 
(Crustacea: Anostraca: Thamnocephalidae) in 
Western Australia, including a description of four 
new species. Hydrobiologia 486, 71-89. 

Timms, B.V. (In press). A key to the fairy shrimps 
(Crustacea: Anostraca) of Australia. Cooperative 
Research Centre Freshwater Ecology Guide, 
Albury. 

Timms, B.V. and Geddes, M.C. (2003). The fairy shrimp 
genus Branchinella Sayce, 1903 (Crustacea: 
Anostraca: Thamnocephalidae) in South Australia 
and the Northern Territory, including descriptions 
of three new species. Transactions Royal Society 
of South Australia 127, 53-68. 

Timms, B.V. and Sanders, P. (2002). Biogeography and 
ecology of Anostraca (Crustacea) in middle Paroo 
catchment of the Australian arid-zone. 
Hydrobiologia 486, 225-238. 

Tommasini, S. and Sabelli, F. S. (1989). Eggshell origin 
and structure in two species of Conchostraca 
(Crustacea, Phyllopoda). Zoomorphology 109, 33- 
37. 

Wolf, E. (1911). Phyllopoda. In Die fauna Sudwest- 
Australiense ergebnisse der Hamburgen sudwest 
australischen Forschungsreise 1905 (Eds W. 
Michaelsen and R. Hartmeyer) (G. Fisher, lena) 
Vol 3, pp 251-276. 



80 Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 



APPENDIX 1 

Locality data for species examined. 

Branchinella affinis: Far North Bulla claypan, 

Rockwell Station, 180 km SW of 

Cunnamulla, Queensland. 28° 52' S, 144° 

56'E, 27-iii-2000. 
Branchinella arborea: Marsilea Pool, Bloodwood 

Station, 130km NW of Bourke, NSW. 29° 

33'S, 144° 52'E, l-vi-1999. 
Branchinella australiensis: Lower Crescent Pool, 

Bloodwood Station, 130 km NW of 

Bourke, NSW, 29° 33'S, 144° 52'E, l-vi- 
1999. 
Branchinella basispina: Homestead Dam, 

Balladonia Station, 220 km E of Norseman, 

WA. 32° 28'S, 123° 52'E, 3-iii-1975. 
Branchinella buchananensis: Gidgee Lake, 

Bloodwood Station, 130 km NE of Bourke, 

NSW, 29° 33'S, 144° 50'E, 28-ix-1998. 
Branchinella budjiti: a claypan on Muella Station, 

128 km NE of Bourke, NSW, 29° 31'S, 

144° 56'E, 6-xii-1999. 
Branchinella campbelli: Muella Lake, Tredega 

Station, 140 km NW of Bourke, NSW, 29° 

31'S, 144° 53'E, 27-vi-1998. 
Branchinella complexidigitata: Lake Arro near 

Eneabba, 300 km N of Perth, WA, 29° 

44'S, 115°10'E, 2-ix-1999. 
Branchinella dubia: pool 89 km from Derby on the 

Gibb R Road, Kimberley, WA, 17° 26' S, 

124° 36'E, 31-1-1985. 
Branchinella frondosa: Steves Pool, Muella Station, 

128 km NW of Bourke, NSW, 29° 33'S, 

144° 55'E, 24-xi-1998. 
Branchinella hattahensis: Mid Kaponyee Lake, 

Currawinya National Park, SW 

Queensland, 28° 50'S, 144° 19'E, 5-vii- 

1997. 
Branchinella kadjikadji: claypan on Kadji Kadji 

Station, 35 km ENE of Morewa, WA, 29° 

08'S, 116°24'E, 14-viii-1998. 
Branchinella lamellata: a claypan near Warburton 

Crossing, Clifton Hills Station, NE South 

Australia, 27° 02'S, 138° 53'E, 5-xii-2000. 
Branchinella longirostris: Warrdagga Rock, via 

Paynes Find, WA, 29° 24'S, 117° 30'E, 26- 

viii-2001. 
Branchinella lyrifera: Turkey claypan, Bloodwood 

Station, 130 km NE of Bourke, NSW, 29° 

34'S, 144° 50'E, l-iv-1999. 
Branchinella nana: Lake Arrow, near Kalgoorlie, 

WA, 30° 32'S, 121° 24'E, 14-V-1995. 



Branchinella nichollsi: Lake Hannan near 

Kalgoorlie, WA, 30° 40'S, 121° 28'E, 17- 

iii-1937. 
Branchinella occidentalis: Freshwater Lake, 

Bloodwood Station, 130 km NW of 

Bourke, 29° 29'S, 144° 50'E, 2-vii-1997 

(Type I); Plover claypan, Bloodwood 

Station, 130 km NW of Bourke, 29° 29'S, 

144° 48'E, 29-vi-1998 (Type H). 
Branchinella pinnata: a blackbox swamp, Tredega 

Station, 140 km NW of Bourke, NSW, 29° 

29'S, 144° 52'E, 30-vi-1999. 
Branchinella probiscida: Darko claypan, 

Currawinya National Park, SW 

Queensland, 28° 52'S, 144° 18'E, 4-xii- 

1999. 
Branchinella simplex: Lake Arrow, near Kalgoorlie, 

WA, 30° 32'S, 121° 24'E, 14-V-1995. 
Branchinella wellardi: Marsilea Pool, Bloodwood 

Station, 130km NW of Bourke, NSW. 29° 

33'S, 144° 52'E, l-vi-1999 
Parartemia contracta: small lake adjacent to Lake 

O'Grady, WA, 30° 25'S, 117° 25'E, ?-viii- 

1978. 
Parartemia cylindrifera: Lake Grace, WA, 33° 06'S, 

18° 24'E, 24-viii-1978. 
Parartemia informis: Lake Ballard, WA, 29° 37' S, 

121° 07'E, 18-viii-1978. 
Parartemia minuta: Lower Bell Lake, Bloodwood 

Station, 130 km NW of Bourke, NSW, 29° 

00'S, 144° 48'E, 28-ix-1998. 
Parartemia zietziana: small lake on 'Flowerfield' 

via Beeac, Victoria, 38° 10'S, 143° 09'E, 

15-viii-1970. 
Streptocephalus queenslandicus: fish rearing ponds, 

Walkamin Research Station, via Atherton, 

Queensland, 17° 08'S, 145° 26'E, ii-1997. 
Streptocephalus n.sp. a: Pine Tree Pool, Bloodwood 

Station, 130 km NE of Bourke, NSW, 29 

29'S, 144 49'E, 2-xii-1999. 
Streptocephalus n.sp.b: Box Hole, 4 km S of 

Homestead, Currawinya National Park, 

Qld., 28 51'S, 144 29'E, l-iv-1999. 
Undescribed new branchiopod genus: Marsilea Pool, 

Bloodwood Station, 130km NW of Bourke, 

NSW. 29° 33'S, 144° 52'E, 24-viii-1998. 



Proc. Linn. Soc. N.S.W., 125, 2004 



81 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 








6 

5 um 


j 




... . ± '^ 


.- - 

It". >■* 






1 


t'*Kr 



Figure 1 . Parartemia contracta cyst. Bar = 50 \im. 

Figure 2. Parartemia contracta cyst shell cross section. Bar = 10 urn. 

Figure 3. Parartemia cylindrifera cyst. Bar = 50 \im. 

Figure 4. Parartemia cylindrifera cyst shell cross section. Bar = lOum. 

Figure 5. Parartemia minuta cyst. Bar = 50 [xm. 

Figure 6. Parartemia minuta cyst shell cross section. Bar = 5 u.m. 



82 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 









!W« 



Figure 7. Parartemia zietziana cyst. Bar = 50 \xm. 

Figure 8. Parartemia zietziana cyst. Bar = 50 [Am. 

Figure 9. Parartemia zietziana cyst shell cross section. Bar = 5 \im. 

Figure 10. Undescribed branchipodid species cyst. Bar = 50 um. 

Figure 11. Undescribed branchipodid species cysts clumped together. Bar = 50 um. 

Figure 12. Undescribed branchipodid species cyst surface. Bar = 10 [jm. 



Proc. Linn. Soc. N.S.W., 125, 2004 



83 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 









Figure 13. Undescribed branchipodid species cyst shell cross section. Bar = 5 urn. 

Figure 14. Streptocephalus queenslandicus cyst. Bar = 50 |j,m. 

Figure 15. Streptocephalus queenslandicus cyst. Bar = 20 |xm. 

Figure 1 6. Streptocephalus queenslandicus cyst shell cross section. Bar = 5 urn. 

Figure 17. Streptocephalus n. sp. a cyst. Bar = 50 um. 

Figure 18. Streptocephalus n. sp. a cyst ridges. Bar = 10 [Am. 



84 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HELL 









Figure 19. Streptocephalus n. sp. a cyst shell cross section. Bar = 5 urn. 

Figure 20. Streptocephalus n. sp. b cyst. Bar = 50 um. 

Figure 21. Streptocephalus n. sp. b cyst ridges. Bar = 10 [un. 

Figure 22. Streptocephalus n. sp. b cyst shell cross section. Bar = 10 [un. 

Figure 23. Branchinella affinis cyst (not quite mature). Bar = 20 \im. 

Figure 24. Branchinella arborea cyst. Bar = 20 [xm. 



Proc. Linn. Soc. N.S.W., 125, 2004 



85 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 













1 




klwkE 






li » • 


HP 1 






Hk *?5 










Figure 25. Branchinella arborea cyst ridge. Bar = 5 urn. 

Figure 26. Branchinella arborea cyst shell cross section. Bar = 5 um. 

Figure 27. Branchinella australiensis cyst. Bar = 20 um. 

Figure 28. Branchinella australiensis cyst ridge structure. Bar = 5 um. 

Figure 29. Branchinella australiensis cyst shell cross section. Bar = 10 um. 

Figure 30. Branchinella basipina cyst. Bar = 50 um. 



86 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 







%€' i " v ' '• 


]CW ^i 




%f^ J£" ^ 
















Figure 31. Branchinella basipina cyst shell cross section. Bar = 10 um. 

Figure 32. Branchinella buchananensis cyst. Bar = 50 um. 

Figure 33. Branchinella buchananensis cyst shell cross section. Bar = 10 um. 

Figure 34. Branchinella budjiti cyst. Bar = 20 um. 

Figure 35. Branchinella budjiti cyst shell cross section. Bar = 5 um. 

Figure 36. Branchinella campbelli cyst. Bar = 20 um. 



Proc. Linn. Soc. N.S.W., 125, 2004 



87 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 









Figure 37. Branchinella campbelli cyst shell cross section. Bar = 5 urn. 
Figure 38. Branchinella complexidigitata cyst. Bar = 50 um. 
Figure 39. Branchinella complexidigitata cyst surface structure. Bar = 10 um. 
Figure 40. Branchinella complexidigitata cyst surface structure. Bar = 10 urn. 
Figure 41. Branchinella complexidigitata cyst shell cross section. Bar = 10 urn. 
Figure 42. Branchinella duhia cyst. Bar = 20 um. 



88 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 






FRONOOSR 


, — _________ 

i 

_rJr* ' * 1 


46 


___r \ 





Figure 43. Branchinella dubia cyst surface. Bar =10 fim. 

Figure 44. Branchinella dubia cyst shell cross section. Bar = 10 [Am. 

Figure 45. Branchinella frondosa cyst. Bar = 20 [Am. 

Figure 46. Branchinella frondosa cyst shell cross section. Bar = 10 [Am. 

Figure 47. Branchinella hattahensis cyst. Bar = 50 [Am. 

Figure 48. Branchinella hattahensis cyst surface structure. Bar = 10 [Am. 



Proc. Linn. Soc. N.S.W., 125, 2004 



89 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 






w* **W : 






Figure 49. Branchinella hattahensis cyst shell cross section with ridge spines. Bar = 10 um. 

Figure 50. Branchinella hattahensis cyst shell cross section. Bar = 10 urn. 

Figure 5 1 . Branchinella kadjikadji cyst. Bar = 50 um. 

Figure 52. Branchinella kadjikadji cyst surface. Bar = 10 um. 

Figure 53. Branchinella kadjikadji cyst shell cross section showing outer sublayer of alveolar layer pulled 

away from inner sublayer. Bar = 5 (Am. 

Figure 54. Branchinella kadjikadji cyst shell fracture showing inner sublayer of alveolar layer. Bar = 2 um. 



90 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 









Figure 55. Branchinella lamellata cyst. Bar = 20 [un. 

Figure 56. Branchinella lamellata cyst shell cross section. Bar = 5 urn. 

Figure 57. Branchinella longirostris cyst. Bar = 50 \xm. 

Figure 58. Branchinella longirostris cyst surface structure. Bar = 10 fun. 

Figure 59. Branchinella longirostris cyst shell cross section. Bar = 10 fun. 

Figure 60. Branchinella lyrifera cyst. Bar = 20 um. 



Proc. Linn. Soc. N.S.W., 125, 2004 



91 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 









Figure 61. Branchinella lyrifera cyst ridges, polar view. Bar = 10 um. 

Figure 62. Branchinella lyrifera cyst ridges, side view. Bar =10 um. 

Figure 63. Branchinella lyrifera cyst shell cross section. Bar = 5 um. 

Figure 64. Branchinella nichollsi cyst. Bar = 50 um. 

Figure 65. Branchinella nichollsi cyst ridge. Bar = 10 um. 

Figure 66. Branchinella nichollsi cyst shell cross section. Bar = 20 \im. 



92 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 









Figure 67. Branchinella occidentalis cyst (type I). Bar = 50 um. 

Figure 68. Branchinella occidentalis cyst (type I) shell cross section. Bar =10 urn. 

Figure 69. Branchinella occidentalis cyst (type II). Bar = 100 um. 

Figure 70. Branchinella occidentalis cyst (type II) surface. Bar = 10 um. 

Figure 71. Branchinella occidentalis cyst (type II) shell cross section. Bar = 20 um. 

Figure 72. Branchinella pinnata cyst. Bar = 20 um. 



Proc. Linn. Soc. N.S.W., 125, 2004 



93 



FAIRY SHRIMP CYST SHELL MORPHOLOGY 









Figure 73. Branchinella pinnata cyst shell cross section. Bar = 10 urn. 

Figure 74. Branchinella probiscida cyst. Bar = 20 urn. 

Figure 75. Branchinella probiscida cyst surface. Bar =10 urn. 

Figure 76. Branchinella probiscida cyst shell cross section. Bar = 5 \im. 

Figure 77. Branchinella simplex cyst. Bar = 20 urn. 

Figure 78. Branchinella simplex cyst shell cross section. Bar = 5 ^m. 



94 



Proc. Linn. Soc. N.S.W., 125, 2004 



B.V. TMMS, W.D. SHEPARD AND R.E. HILL 






Figure 79. Branchinella wellardi cyst. Bar = 20 urn. 

Figure 80. Branchinella wellardi cyst ridges. Bar =10 |un. 

Figure 81. Branchinella wellardi cyst shell cross section. Bar = 10 um. 



Proc. Linn. Soc. N.S.W., 125, 2004 



95 



96 



The Yule Island Fauna and the Origin of Tropical Northern 
Australian Echinoid (Echinodermata) Faunas 

I.D. LlNDLEY 

Department of Geology, Australian National University, Canberra, A.C.T. 0200. 
(lindley@geology.anu.edu.au) 



Lindley, I.D. (2004). The Yule Island Fauna and the Origin of Tropical Northern Australian Echinoid 
(Echinodermata) Faunas. Proceedings of the Linnean Society of New South Wales 125, 97-109. 

Systematic description of the rich Lower Pliocene echinoid fauna from Yule Island, Papua New 
Guinea, and recently available palaeogeographic data from onshore Papua, the Gulf of Papua and Torres 
Strait have provided new insights into the origins of the extant tropical northern Australian echinoid fauna. 
Previous studies of echinoderm origins, hindered by a lack of fossil evidence, concluded that tropical northern 
Australian echinoderms were derived predominantly by Recent migrations from East Indian and West Pacific 
stocks. However, 47 per cent of species from the Yule Island fauna are extant in northern Australian waters, 
indicating that present faunistic patterns were to a large extent, in-place by at least the Lower Pliocene. 
Palaeogeographic evidence supports the earlier observations of H. Barraclough Fell, that migration of echinoid 
stock into (and out of) eastern New Guinea and tropical northern Australia probably occurred during the 
Lower to Middle Miocene, when widespread tropical to sub-tropical reef development occurred across a 
5600 km belt from SE Asia through New Guinea and into the SW Pacific as far as Fiji. This favourable 
pathway for exchange between echinoid stocks disappeared during the Upper Miocene, when the onset of 
tectonic instability throughout the region, and the establishment of a discontinuous volcanic arc, resulted in 
influx of terrigenous sediments and may have caused the death of the reef complex. This pattern of 
sedimentation has persisted to the present. 

Manuscript received 11 April 2003, accepted for publication 20 August 2003. 

KETWORDS: Biogeography, East Indies, Echinoidea, Great Barrier Reef, Palaeogeography, Papua New 
Guinea, Pliocene, Queensland. 



the only tropical Miocene macro-invertebrate fauna 

INTRODUCTION in Australia (McNamara and Kendrick 1994). The 

diverse Mio-Pliocene echinoid fauna described by 

Echinoderms are one of the most intensively Jeannet and R. Martin (1937) from Java, about 1 000 

studied shallow-water faunas of tropical northern km WNW of the northern GBR, provides closer 

Australia (Endean 1982). Their specific composition (geological) time links with the region than does the 

and general distribution in the region is well known Barrow Island fauna. However, caution is necessary 

because of the work of A.H.Clark (1908, 1911a, 1915, in reviewing Jeannet and R. Martin's (1937) fauna, 

1915-1967), H.L. Clark (1907, 1909, 1915, 1921, with the validity of some species identifications in need 

1925, 1926, 1928, 1932, 1938, 1946), Endean (1953, of re-evaluation and name changes required, in 

1956, 1957, 1961, 1965), A.M. Clark (1970), A.M. accordance with present taxonomic nomenclature. The 

Clark and Rowe (1971), and Gibbs et al. (1976). rich Pliocene Yule Island, Papua New Guinea, echinoid 

However, studies of the origins of these echinoderm fauna comprising 19 species, is located less than 300 

faunas have been hindered by the lack of outcropping km from the northern end of the GBR, and represents 

marine Tertiary sequences with macro-invertebrate the nearest Tertiary echinoid fauna to the GBR region 

faunas (Endean 1957). McNamara and Kendrick (Lindley 2003a-c; Fig. 1). This paper examines the 

(1994) described the Middle Miocene echinoid fauna importance of the Yule Island fauna with respect to 

from the Poivre Formation on Barrow Island, off the the origins of extant echinoid faunas of tropical 

Western Australian coast, about 1 500 km WSW of northern Australia. 

the northern Great Barrier Reef (GBR). The Poivre The taxonomic classification used herein 

Formation represents the northern-most exposure of f n ows that of Fell (1966), Fell and Pawson (1966), 

Miocene marine deposits in Australia, and contains Durham (1966) and Fischer (1966). 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 




AUSTRALIA 



Figure 1. Locality map showing proximity of Yule Island to tropical northern Australia. 



FAUNAL PROVINCES AND MAINLAND AND 
REEF ECHINODERM FAUNAS 

Tropical northern Australian seas embrace 
two faunistic provinces, namely, the Tropical 
Australian Province and the Solanderian Province 
(Endean 1957). 

The Solanderian Province (Whitley 1932) is 
restricted to the fauna of the GBR, and includes 
echinoderms which Endean (1957) designated as 'reef 
species (Fig. 2). These echinoderms are considered to 
have strong affinities with West Pacific stocks, with 
their pelagic larval stages transported onto the GBR 
by the Pacific south equatorial current Endean (1957). 
Endean (1957) considered that the lack of reef 
structures to the west of Torres Strait has proved a 
major barrier to the migration of reef stocks into the 
Solanderian Province. 

The Tropical Australian Province of Endean 
(1957) incorporates the Banksian Province of Whitley 
(1932) and the Dampierian Province of Hedley (1904, 
1926) and extends from Geraldton, on the Western 



Australian coast, to Wide Bay (26°S) on the southern 
Queensland coast (Fig. 2). As originally proposed, the 
Banksian Province included the marine faunas of 
coastal districts of Queensland, distinct from those of 
the GBR, and the Dampierian Province along the 
Western Australian coast. However, Endean (1957) 
concluded that Torres Strait does not present a major 
biogeographical boundary separating the Dampierian 
and Banksian Provinces, and that they should be 
considered one single faunistic unit. The Tropical 
Australian Province contains an echinoderm fauna 
designated by Endean (1957) as 'mainland' species, 
typically found in habitats dominated by terrigenous 
sediments. Mainland echinoderms have strong 
affinities with East Indies stocks, with the principal 
exchange route being via Torres Strait and the Arafura 
Sea (Endean 1957). He noted that for Queensland 
waters, there is little intermingling between reef and 
mainland stocks. 

The East Indies Province lies to the north of 
Australia (Fig. 2). H.L. Clark (1946) defined the 
geographic limits of this faunal province, to include 
the sea and its islands between 90° and 155°E, with a 



98 



Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 




Figure 2. Australia and New Guinea showing the distribution of 
biogeographical zones mentioned in the text. 



H. Barraclough Fell 

Fell (1953) also 
concluded that there had 
been a mainly southward 
migration along the Indo- 
Malayan archipelago, but 
noted that northward 
movements of Australasian 
echinoderms into the Indo- 
Pacific could be detected 
from the Miocene onward. 
He believed that the 
archipelago, or its Tertiary 
equivalents, provided the 
shallow-water migration 
route, both into and out of 
Australia. Fell (1953) noted 
that profound changes in 
fossil Australian echinoid 
faunas occurred during the 
Upper Miocene, an event he 
attributed to cooling climate. 



southern limit coinciding with the northern limit of 
what is now know as the Tropical Australian Province. 
At its eastern extent, it includes the island of New 
Guinea with the eastern boundary following longitude 
155° between the Bismarck Archipelago and the 
Solomon Islands to 5°N, where it turns west to 130°E. 



PREVIOUS BIOGEOGRAPHICAL STUDIES 

Austin Hobart Clark and Hubert Lyman Clark 

Much of our knowledge of shallow-water 
Australian echinoderms is based on the intensive work 
of A.H. Clark and H.L. Clark. A.H. Clark (1911b) 
concluded that 'The crinoids of Australia have come 
from the north, from the great East Indian 
Archipelago' . H.L. Clark (1946) extended A.H. Clark's 
observations to other echinoderm groups, and 
concluded that the 'evidence is overwhelming that 
Australian echinoderms have come southward from 
the East Indian area, either around the eastern end of 
New Guinea or across the Timor Sea'. H.L. Clark 
(1921, 1946) thought that the extant Queensland fauna 
postdated the 'depression of land areas east of New 
Guinea which led to the connection of the Coral Sea 
with the western Pacific and the East Indian region'. 
The Dampierian fauna had migrated from farther west 
and was well established when formation of the Torres 
Strait made mingling of the two faunas possible. 



Robert Endean 

Endean (1957) saw several obstacles to H.L. 
Clark's (1946) echinoderm migration patterns around 
New Guinea. Firstly, he believed there was little 
likelihood of any significant exchange of mainland 
species via the northern shores of New Guinea. Deep 
waters are present immediately offshore of the north 
coast of the island, and the large Sepik and 
Mamberamo Rivers, by contributing large volumes of 
freshwater and silt, acted as natural barriers to the 
spread of littoral echinoderms. He argued that in 
eastern New Guinea there was a lack of suitable 
habitats for mainland echinoderms, with the Papua- 
Louisiade Barrier Reef, extending along the 
southeastern coast of New Guinea and to the east of 
the island, representing an additional barrier to the 
migration of mainland species. Endean (1957) 
considered that the Fly River, which empties large 
amounts of silt and freshwater into the Gulf of Papua, 
acted as an ecological barrier to the spread of reef 
species to the west of Torres Strait. 

Endean (1957) concluded that extant tropical 
northern Australian echinoderm faunas were derived 
predominantly from Recent migrations of East Indian 
and West Pacific stocks. With the Queensland 
mainland species having strong affinities with those 
of the East Indies, the principal exchange route of these 
forms was via the Torres Strait and the Arafura Sea, 
the deepwater of the Coral Sea serving as a barrier to 
the spread of mainland species to the West Pacific. 



Proc. Linn. Soc. N.S.W., 125, 2004 



99 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 



Endean (1957) believed that reef species have strong 
ties with those of the West Pacific. Given that very 
few of the reef species which occur outside of 
Australian waters are present in waters west of Torres 
Strait, he concluded that there was insignificant 
exchange of reef species in Queensland waters with 
those of the East Indies by way of the Torres Strait. 
Rather, the main route of exchange of Queensland and 
West Pacific echinoderms must have been by way of 
either the south coast of eastern New Guinea or the 
Coral Sea. However, since eastern New Guinea and 
the GBR are separated by deep water from West Pacific 
localities, Endean (1957) believed the most likely route 
of exchange of Queensland's reef species and West 
Pacific stocks must have been via pelagic larval stages 
swept into the Coral Sea by the unidirectional Pacific 
south equatorial current. 



ANALYSIS OF THE YULE ISLAND ECHINOID 

FAUNA 

The echinoid faunas from the Lower Pliocene 
of Yule Island, Middle Miocene of Barrow Island and 
the Mio-Pliocene of Java each contain mixed 
assemblages of clypeasteroid, regular and spatangoid 
echinoids, representative of very different 
palaeoecologies. Because none of these faunas is 
dominated by an echinoid group unique to a particular 
palaeoecology, valid comparisons can be made 
between these fossil faunas. Using a similar argument, 
these comparisons are extended to the extant faunas 
of tropical northern Australia. 

Of the 19 echinoid species described from 
Yule Island, a group of seven species is not recorded 
from either the Barrow Island and Javanese faunas or 
the extant echinoid fauna of tropical northern Australia 
(Table 1 - located after the reference list). These species 
include Schizechinus cf. tuberculatus (Pomel), 
Phyllacanthus sp., Laganum depressum sinaiticum 
Fraas 1867, and L. depressum delicatum Mazzetti 
1894, Palaeostoma kairukuensis Lindley 2003c, 
Maretia cordata Mortensen 1948, and Schizaster 
(Schizaster) alphonsei Lindley 2003c. Only two of 
these echinoids are endemic, viz. S.(S.) alphonsei and 
P. kairukuensis. Laganum depressum sinaiticum, and 
L. depressum delicatum are notable in that they are 
otherwise only recorded extant in the western Indian 
Ocean (Persian Gulf). Similar observations of western 
Indian Ocean affinities have been made amongst fossil 
reef corals from the nearby Upper Pliocene-Lower 
Pleistocene Era Beds, northwest of Yule Island, notably 
the faviid coral Parasimplastrea simplicitexta 
Umbgrove 1942 (Veron and Kelley 1988). 



From an analysis of the remaining 12 species, 
nine occur in tropical waters of northern Australia 
either in mainland or reef stocks. These species include 
Cyrtechinus verruculatus (Liitken), Prionocidaris 
verticillata (Lamarck 1816), Parasalenia poehli 
Pfeffer 1887, Clypeaster humilis (Leske), C. latissimus 
(Lamarck), C. reticulatus (Linnaeus 1758), Laganum 
depressum Lesson in L. Agassiz 1841, L. decagonale 
(de Blainville 1827) and Maretia planulata (Lamarck). 
Nine species are known as fossil in the Mio-Pliocene 
of Java, viz. Prionocidaris verticillata, Phyllacanthus 
imperialis var. javana K. Martin 1885, Temnotrema 
macleayana (Tenison-Woods 1878), L. depressum, L. 
decagonale, C. reticulatus, C. humilis, M. planulata 
and Eupatagus (Eupatagus) pulchellus (Herklots). The 
presence of these species as fossil in the Mio-Pliocene 
of Java, and the observed low levels of species 
endemism, suggests that the two populations were 
well-connected prior to and during the Pliocene. By 
contrast, the Yule Island fauna, as with the Javanese 
fauna, does not have any species in common with the 
Middle Miocene fauna of Barrow Island (Table 1). 

Significantly, 47 per cent of echinoid taxa 
(nine species) from the Yule Island fauna, now inhabit 
northern Australian waters. Clearly, this observation 
conflicts with Endean' s (1957) proposal that tropical 
Australian echinoderms were derived from Recent 
migrations from East Indian and West Pacific stocks. 
Of these nine species, four can be included as mainland 
stock, viz. C. verruculatus, C. reticulatus, C. humilis 
and C. latissimus; three are of reef stock, viz. L. 
decagonale, P. verticillata and P. poehli; and two 
species occur in both reef and mainland waters viz. L. 
depressum and M. planulata. Endean (1957) noted a 
similar, but limited intermingling of echinoderm stock, 
including the echinoids M. planulata and Salmacis 
sphaeroides, in waters surrounding the Low Isles, a 
coral-dominated locality relatively close to the 
mainland (about 1 1 km distant), north of Cairns. He 
regarded these species as having 'strayed into the coral 
environment'. Such a near-shore intermingling is 
envisaged during the Pliocene of Yule Island (Fig. 3). 



MIO-PLIOCENE PALAEOGEOGRAPHY OF THE 
GULF OF PAPUA AND ADJACENT AREAS 

Advances in the knowledge of the geological 
evolution of onshore Papua and the western Coral Sea 
have emerged with the synthesis of extensive oil well 
and outcrop data gathered during hydrocarbon 
exploration of the past 25 years (Home et al. 1990; 
Carman 1993; Struckmeyer et al. 1993 amongst 
others). These advances have permitted a detailed 



100 



Proc. Linn. Soc. N.S.W., 125, 2004 



ID. LINDLEY 



r 

o 




CD 


CD 




H 


CD 










■D 






CD 


CO 




CO 


U) 




o 


CO 




-C 


^ 


CO 


+2 


CO 


r 


o 


h 


o 


c 
CO 


o 


Q. 


o 


o 


« 


> 


m 


a 



3dlN3COd3a 
QNV1XOIU1S 



i -a 

OX) o 

£ -S 
?«« 

DC cs 

e © 

]e 

« je 
.2 © 

DO C ^ 

^- S o\ 

•w ™ ©> 

"« DC w 

vi « g 

£ I u 

at ++ S 

* - t§ 

cS g a» 

3 fa a> 

&£* 

21 E 

g .fa a 

^ DC W 

&''« 5 

s- — © 

DC W ** 

2 S <« 

DC 5 

® e P 

05 J" {y 

^ ,2 a 

o <u — 

. CU 

.2 2 § 

2 fc o 

35S 

DC ^ 3 

E.SO 



understanding of the development of Mio-Pliocene 
sedimentary facies and the region's tectonic history. 
Tectonic stability during the Lower to Middle 
Miocene was associated with the development of an 
extensive tropical to sub-tropical platform carbonate 
reef complex, not only in the Papuan Basin, but in a 
region extending from SE Asia to Fiji, a distance of 
some 5 600 km, and including northern Australia 
(Coleman and Packham 1976; Home et al. 1990; 
McNamara and Kendrick 1994). During the late 
Miocene, the onset of tectonic compression along the 



edge of the Pacific Plate across much of the region 
resulted in major uplift and the establishment of a 
discontinuous volcanic arc. In New Guinea, as 
elsewhere throughout the region, the influx of 
terrigenous sediments, to the north and south of an 
uplifting central cordillera, resulted in the death of the 
Miocene reef complex (Home et al. 1990; Carman 
1993; Struckmeyer et al. 1993). The western end of 
the Gulf of Papua became emergent during this time 
and sedimentation was predominantly on a fluvial/ 
alluvial plain adjacent to a volcanic province (Carman 



Proc. Linn. Soc. N.S.W., 125, 2004 



101 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 



1993; Fig. 3). South of a fluctuating Pliocene shoreline, 
poorly sorted deltaic/pro-deltaic sediments were 
deposited across a greater part of the rapidly subsiding 
Gulf of Papua (Home et al. 1990; Carman 1993). More 
than 4 000 m of sediment were deposited in the Vailala- 
Purari depocentre, near the mouth of the Purari River 
(Carman 1993), indicative of very rapid subsidence. 
Subsidence and the large quantities of clastic sediment 
were deterrents to coral growth across a large part of 
the region. However, biohermal reef limestone 
(foraminiferal zone c. N20: Lower Pliocene/Upper 
Pliocene boundary), surrounded by interpreted 
argillaceous micritic limestone, penetrated in the 
Anchor Cay 1 Well, only 250 km WSW of Yule Island, 
is associated with reefal precursors of the present day 
GBR which existed farther south (Carman 1993). 
These patterns of sedimentation have continued to the 
present day (Home et al. 1990). 



ORIGIN OF TROPICAL AUSTRALIAN 
ECHTNOID FAUNAS 

Preamble 

Migration of adult littoral echinoderms may 
occur via connected shallow-water, typically near- 
shore habitats. Deep water, a lack of interconnected 
reefal structures, and large influxes of freshwater and 
silt poured out from large coastal rivers, all act as 
ecological barriers to their spread (Endean 1957). 
However, for echinoderms possessing prolonged larval 
stages, transport of pelagic young by currents, may 
distribute species across deep water (Endean 1957; 
Nichols 1969). 

Lower to Middle Miocene origin of the Yule Island 
fauna 

The Yule Island echinoid fauna comprises 
representatives of reef and mainland species and 
species common to both reef and mainland faunas. 
Comparisons show that, with the relatively low level 
of species endemism evident in the Yule Island 
echinoid fauna, prior to the Lower Pliocene the fauna 
was well-connected with at least the population in Java, 
having not developed in isolation. Palaeogeographic 
evidence indicates that an excellent interconnected 
exchange route for echinoderms existed during the 
Lower to Middle Miocene when an extensive tropical 
to sub-tropical reef complex existed across a 5 600 
km belt extending from SE Asia through New Guinea 
to Fiji. Both reef and mainland species migrated along 
this reef complex. 

Endean' s (1957) West Pacific influence on 
extant reef echinoid populations of tropical northern 
Australia may be more apparent than real. For all 



echinoids, reef and mainland, only one, Rhynobrissus 
hemiasteroid.es Agassiz, is endemic to the West Pacific. 
Eleven species occur in the Indian Ocean and East 
Indies, and 21 are widely distributed throughout the 
Indo-Pacific. The origins of the reef stock at Yule 
Island, with its apparent 'West Pacific affinity', is 
readily accounted for by the interconnected pathway 
afforded by the 5 600 km long Lower to Middle 
Miocene reef complex that extended well into the SW 
Pacific. 

Fell (1953) noted that a great extinction of 
the Australasian echinoid stocks occurred during the 
late Miocene, an event he and Fleming (1949) 
attributed to a cooling climate. The cooling event 
coincided with the previously noted retreat of 
widespread platform carbonate deposition from SE 
Asia and the SW Pacific. However, the demise of the 
Miocene carbonate platform in the western Coral Sea 
was accompanied by a resurgence of tectonic instability 
and associated volcanism. This tectonic instability is 
evident in the post-Miocene geological record that 
succeeds carbonate deposition throughout the region 
(Coleman and Packham 1976) and it is possible that 
an increase in volcanic activity, and in-turn 
atmospheric volcanic aerosols, was responsible for the 
late Miocene cooling climate. 

The late Miocene extinction saw the 
disappearance of, for example, the warm-water 
echinoid genera Schizaster L. Agassiz 1836 and 
Phyllacanthus Brandt 1835 from New Zealand (Fell 
1953). Fell (1953) considered that the impact of the 
extinction event in low latitudes was reduced. A 
comparison of the Yule Island fossil fauna with low 
latitude Miocene faunas from India, Java and Fiji 
supports this view. Eight species from the Yule Island 
fauna, P. imperialis var. javana, P. verticillata, T. 
macleayana, L. decagonale, L. depressum, C. 
reticulatus, C. humilis and E.(E.) pulchellus are known 
from the Miocene faunas of these regions (Jeannet and 
R. Martin 1937; Mortensen 1948; Lindley 2001, 
2003a-c). 

Origin of Tertiary and extant northern Australian 
echinoid faunas 

As previously noted, mainland and reef 
echinoid populations are believed to have established 
themselves in the region to the north of Australia and 
adjacent areas during the Lower to Middle Miocene 
when platform carbonate sedimentation prevailed 
throughout much of SE Asia and the SW Pacific. Fell 
(1953) proposed a broadly similar outline for 
Australasian echinoid origins. 

The Middle Miocene echinoid fauna from the 
Poivre Formation on Barrow Island is a far from 
complete representation of the original fauna 



102 



Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



(McNamara and Kendrick 1994). The fauna (at a 
generic level) has much in common with the Miocene 
faunas of Java and India (McNamara and Kendrick 
1994), but its relative geographic isolation during the 
Miocene, 15-20°S of Yule Island and Java, may 
explain the faunal mismatch at species level. Barrow 
Island during the early Miocene was located about 
40°S, Yule Island 25 °S and eastern Indonesia 20°S 
(Veevers et al. 1991: Fig. 12). McNamara and 
Kendrick (1994) noted that, at a generic level, the 
echinoid fauna has strong affinities with modern 
communities of the region. 

Endean (1957) considered that the extant NW 
Australian echinoderm fauna, dominated by mainland 
species, has strong affinities with East Indies stocks, 
but exhibits a high degree of endemism. The deep water 
of the Timor and Arafura Seas appears to have served 
as a barrier to any significant post-Miocene exchange 
between the East Indies and NW Australian mainland 
faunas, and exchange with Queensland mainland 
faunas has only been possible since the opening of 
Torres Strait. The strait was emergent during at least 
the late Miocene-early Pliocene (Fig. 3). Ekman (1953) 
noted that the average age of echinoid species was at 
most 4-6 million years, and the origin of many endemic 
species in the NW Australian mainland fauna may have 
resulted from the long period of geographic isolation 
following the demise of the Lower to Middle Miocene 
(10-20 Ma) reef. 

The Lower Pliocene coral growth at Yule 
Island (foraminiferal zones N18/N19-N20: 4-6 Ma) 
was part of a chain of interconnected reefs extending 
NW along the northern margin of the Gulf of Papua, 
remnants of the formerly extensive Miocene reef 
complex (Fig. 3). Reef species were confined to these 
coral structures, which were in-turn flanked by 
terrigenous sediments, a habitat dominated by 
mainland species. In the southern Gulf of Papua, reef 
precursors to the GBR did not appear until 3-4 Ma 
(Lower Pliocene/Upper Pliocene boundary: 
foraminiferal zone c. N20: Carman 1993; Haig 1996). 
These precursor reefs were situated only 250 km WSW 
of Yule Island and extended farther south (Carman 
1993), indicating that, although large quantities of 
terrigenous sediment at this time were deterrents to 
coral growth, reefs were able to establish themselves 
in parts of the Gulf of Papua. Of the nine echinoid 
species (47 per cent) of the Yule Island fauna that are 
recorded extant in northern Australian waters, five are 
recorded on the GBR, suggestive of ties between both 
faunas. With major barriers to echinoderm migration 
existing to the west (Torres Strait was emergent) and 
along northern New Guinea (deep waters and 
terrigenous sedimentation), it is likely that species 



exchange occurred from existing Pliocene Gulf of 
Papua echinoid stocks to proto-GBR structures and 
farther south. 



ACKNOWLEDGMENTS 

The author kindly acknowledges Prof. K.S.W. 
Campbell for his constructive review of and improvements 
to the manuscript. The comments of anonymous referees 
and Dr M.L. Augee also improved the manuscript. 

REFERENCES 

Carman, G.J. (1993). Palaeogeography of the Coral Sea, 
Darai and Foreland megasequences in the 
eastern Papuan Basin. In 'Petroleum Exploration 
and Development in Papua New Guinea: 
Proceedings of the Second PNG Petroleum 
Convention, Port Moresby' (Eds. G.J. Carman 
and Z, Carman) pp. 291-309. (PNG Chamber of 
Mines and Petroleum: Port Moresby). 

Clark, A.H. (1908). New genera and species of crinoids. 
Proceedings of the Biological Society of 
Washington 21, 219-231. 

Clark, A.H. (1911a). The recent crinoids of Australia. 

Memoirs of the Australian Museum 4, 706-804. 

Clark, A.H. (1911b). The comparative age of the recent 

crinoid faunas. American Journal of Science 32, 
127-132. 

Clark, A.H. (1915). The distribution of the recent crinoids 
on the coasts of Australia. Internationale Revue 
der Gesamten Hydrobiologie und Hydrographie 
7, 222-234. 

Clark, A.H. (1915-1967). A monograph of the existing 
crinoids. Bulletin of the United States National 
Museum 82: 1-406 (1915); 1-795 (1921); 1-816 
(1931); 1-603 (1941); 1-473 (1947); 1-383 
(1950); 1-860 (1967). 

Clark, A.M. (1970). The Swain Reefs expedition: 

Crinoidea. Records of the Australian Museum 
29, 391-406. 

Clark, A.M. and Rowe, F.W.E. (1971). Monograph of 
shallow-water Indo-West Pacific echinoderms. 
British Museum (Natural History) Publication 
690, 238pp. 

Clark, H.L. (1907). The cidaridae. Bulletin of the Museum 
of Comparative Zoology, Harvard 51, 165-228. 

Clark, H.L. (1909). Notes on some Australian and Indo- 
Pacific echinoderms. Bulletin of the Museum of 
Comparative Zoology, Harvard 52, 107-137. 

Clark, H.L. (1915). The comatulids of Torres Strait with 
special reference to their habits and reactions. 
Carnegie Institution of Washington, Publication 
212, 97-125. 

Clark, H.L. (1921). The echinoderm fauna of Torres 

Strait: its composition and its origin. Carnegie 
Institution of Washington, Publication 214, 
223pp. 



Proc. Linn. Soc. N.S.W., 125, 2004 



103 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 



Clark, H.L. (1925). A catalogue of the recent sea-urchins 
(Echinoidea) in the collection of the British 
Museum (Natural History). British Museum 
(Natural History), 250pp. 

Clark, H.L. (1926). Notes on a collection of echinoderms 
from the Australian Museum. Records of the 
Australian Museum 15, 183-192. 

Clark, H.L. (1928). The sea-lilies, sea-stars, brittle stars 
and sea urchins of the South Australian 
Museum. Records of the South Australian 
Museum 3(4), 361-482. 

Clark, H.L. (1932). Echinodermata (other than 

Asteroidea). Scientific Report of the Great 
■ Barrier Reef Expedition 4, 197-239. 

Clark, H.L. (1938). Echinoderms from Australia: An 

account of collections made in 1929 and 1932. 
Memoirs of the Harvard Museum of 
Comparative Zoology 55, 596pp. 

Clark, H.L. (1946). The Echinoderm Fauna of Australia: 
Its Composition and Its Origin. Carnegie 
Institution of Washington, Publication 566, 
567pp. 

Coleman, P.J. and Packham, G.H. (1976). The Melanesian 
borderlands and India-Pacific plates' boundary. 
Earth Science Reviews 12, 197-233. 

Durham, J.W. (1966). Clypeasteroids. In 'Treatise on 
Invertebrate Paleontology, Part U, 
Echinodermata 3' (Ed. R.C Moore) pp. U450- 
U491. (Geological Society of America and 
University of Kansas Press: Lawrence). 

Ekman, L. (1953). 'Zoogeography of the Sea'. (William 
Clowes and Sons: London). 

Endean, R. (1953). Queensland faunistic records. 3. 

Echinodermata (excluding Crinoidea). Papers of 
the Department of Zoology, University of 
Queensland 1, 53-60. 

Endean, R. (1956). Queensland faunistic records. 4. 

Further records of Echinodermata (excluding 
Crinoidea). Papers of the Department of 
Zoology, University of Queensland 1, 123-140. 

Endean, R. (1957). The biogeography of Queensland's 
shallow-water echinoderm fauna {excluding 
Crinoidea), with a rearrangement of the faunistic 
provinces of tropical Australia. Australian 
Journal of Marine and Freshwater Research 8, 
233-273. 

Endean, R. (1961). Queensland faunistic records. 7. 

Additional records of Echinodermata (excluding 
Crinoidea). Papers of the Department of 
Zoology, University of Queensland 1, 289-298. 

Endean, R. (1965). Queensland faunistic records. 8. 

Further records of Echinodermata (excluding 
Crinoidea). Papers of the Department of 
Zoology, University of Queensland 2, 229-235. 

Endean, R. (1982). 'Australia's Great Barrier Reef. 
(University of Queensland Press: St. Lucia). 

Fell, H.B. (1953). The origin and migrations of 

Australasian echinoderm faunas since the 
Mesozoic. Transactions of the Royal Society of 
New Zealand 81, 245-255. 



Fell, H.B. (1966). Cidaroids. In 'Treatise on Invertebrate 
Paleontology, Part U, Echinodermata 3' (Ed. 
R.C. Moore) pp. U312-U340. (Geological 
Society of America and University of Kansas 
Press: Lawrence). 

Fell, H.B. and Pawson, D.L. (1966). Echinacea. In 

'Treatise on Invertebrate Paleontology, Part U, 
Echinodermata 3' (Ed. R.C. Moore) pp. U367- 
U440. (Geological Society of America and 
University of Kansas Press: Lawrence). 

Fischer, A.G. (1966). Spatangoids. In 'Treatise on 
Invertebrate Paleontology, Part U, 
Echinodermata 3' (Ed. R.C. Moore) pp. U543- 
U628. (Geological Society of America and 
University of Kansas Press: Lawrence). 

Fleming, CA. (1949). The geological history of New 
Zealand. Tuatara 2(2), 72-90. 

Gibbs, P.E., Clark, A.M. and Clark, CM. (1976). 

Echinoderms from the northern region of the 
Great Barrier Reef, Australia. Bulletin of the 
British Museum (Natural History) Zoology 
30(4), 101-144. 

Haig, D.W. (1996). Late Neogene bathyal depocentres in 
mainland Papua New Guinea. In 'Petroleum 
Exploration, Development and Production in 
Papua New Guinea: Proceedings of the Third 
PNG Petroleum Convention, Port Moresby' 
(Ed. P.G. Buchanan) pp. 313-327. (PNG 
Chamber of Mines and Petroleum: Port 
Moresby). 

Hedley, C. (1904). The effect of the Bassian Isthmus upon 
the existing marine fauna: a study in ancient 
geography. Proceedings of the Linnean Society 
of New South Wales 28, 876-883. 

Hedley, C. (1926). Zoogeography. In 'Australian 

Encyclopedia' Vol. 2, pp. 743. (Angus and 
Robertson: Sydney). 

Home, P.C, Dalton, D.G. and Brannan, J. (1990). 

Geological evolution of the western Papuan 
Basin. In 'Petroleum Exploration in Papua New 
Guinea: Proceedings of the First PNG Petroleum 
Convention, Port Moresby' (Eds. G.J. Carman 
and Z. Carman) pp. 107-1 17. (PNG Chamber of 
Mines and Petroleum: Port Moresby). 

Jeannet, A. and Martin, R. (1937). Ueber Neozoische 

Echinoidea aus dem Niederlaendisch-Indischen 
Archipel. Leidsche geologische mededeelingen 
8(2), 215-308. 

Lindley, I.D. (2001). Tertiary Echinoids from Papua New 
Guinea. Proceedings of the Linnean Society of 
New South Wales 123, 119-139. 

Lindley, I.D. (2003a). Echinoids of the Kairuku 

Formation (Lower Pliocene), Yule Island, Papua 
New Guinea: Clypeasteroida. Proceedings of the 
Linnean Society of New South Wales 124, 125- 
136. 

Lindley, I.D. (2003b). Echinoids of the Kairuku 

Formation (Lower Pliocene), Yule Island, Papua 
New Guinea: Regularia. Proceedings of the 
Linnean Society of New South Wales 124, 137- 
151. 



104 



Proc. Linn. Soc. N.S.W., 125, 2004 



ID. LINDLEY 



Lindley, I.D. (2003c). Echinoids of the Kairuku 

Formation (Lower Pliocene), Yule Island, Papua 
New Guinea: Spatangoida. Proceedings of the 
Linnean Society of New South Wales 124, 153- 
162. 

McNamara, K.J. and Kendrick, G.W. (1994). Cenozoic 
molluscs and echinoids of Barrow Island, 
Western Australia. Records of the Western 
Australian Museum Supplement No. 51, 50pp. 

McNamara, K.J. and Philip, G.M. (1980). Living 

Australian schizasterid echinoids. Proceedings 
of the Linnean Society of New South Wales 104, 
127-146. 

Mortensen, T. (1943). A Monograph of the Echinoidea 
3(2), Camarodonta 1. (C.A. Reitzel, 
Copenhagen). 553pp. 

Nichols, D. (1969). 'Echinoderms'. (Hutchinson 
University Library: London). 192pp. 

Philip, G.M. (1963). The Tertiary echinoids of 

southeastern Australia. 1 Introduction and 
Cidaridae (1). Proceedings of the Royal Society 
of Victoria 76, 181-226. 

Struckmeyer, H.I.M., Yeung, M. and Pigram, C.J. (1993). 
Mesozoic to Cainozoic plate tectonics and 
palaeogeographic evolution of the New Guinea 
region. In 'Petroleum Exploration and 
Development in Papua New Guinea: 
Proceedings of the Second PNG Petroleum 
Convention, Port Moresby' (Eds. GJ. Carman 
and Z. Carman) pp. 262-290. (PNG Chamber of 
Mines and Petroleum: Port Moresby). 

Veevers, J.J. Powell, C.McA. and Roots, S.R. (1991). 

Review of seafloor spreading around Australia. 
1 . Synthesis of the patterns of spreading. 
Australian Journal of Earth Sciences 38, 373- 
389. 

Veron, J.E.N. and Kelley, R. (1988). Species stability in 
reef corals of Papua New Guinea and the Indo 
Pacific. Association of Australasian 
Palaeontologists, Memoir 6, 69pp. 

Whitley, G. (1932). Marine zoogeographical regions of 
Australia. Australian Naturalist 8, 166-167. 



Proc. Linn. Soc. N.S.W., 125, 2004 105 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 



TABLE 1 : Distribution of selected shallow-water tropical echinoid groups in the East Indies and Australia. 
Compiled from Endean (1957); A.M. Clark and Rowe (1971); Gibbs et al. (1976); Jeannet and R. Martin 
(1937); Philip (1963); Mortensen (1943); H.L. Clark (1946); Lindley (2001, 2003a,b,c); McNamara and 

Kendrick (1994); and McNamara and Philip (1980). 


Species 


Java, Mio- 
Pliocene 


Yule Is, 

Lower 

Pliocene 


Barrow Is, 

Middle 

Miocene 


Australian 

'mainland' 

species 


Australian 

'reef 

species 


CIDARIDAE 












Phyllacanthus dubius 


X 










Phyllacanthus dubius var. 
sundaica 


X 










Phyllacanthus imperialis 


X 






X 


X 


Phyllacanthus imperialis var. 
iavana 


X 


X 








Phyllacanthus sp. 




X 








Phyllacanthus cf. clarkeii 






X 






Prionocidaris bispinosa 








X 


X 


Prionocidaris verticillata 


X 


X 






X 


Prionocidaris baculosa 


X 










Prionocidaris baculosa var. 
annulifera 


X 










Stylocidaris reini 


X 










Cidaris mertoni 


X 










Cidaris aculeata 


X 










Cidaris sp. 


X 










Chondrocidaris sundaica 


X 










Eucidaris sp. 






X 






Goniocidaris cf. murrayensis 






X 


r 
















DIADEMATIDAE 












Diadema setosum 








X 


X 


Diadema savignyi 










X 


Echinothrix calamaris 










X 


Echinothrix diadema 










X 


Astropyga radiata 








X 
















PARASALENIIDAE 












Parasalenia poehli 




X 






X 


Parasalenia gratiosa 








X 


X 


Parasalenia sp. 










X 














TOXOPNEUST1DAE 












Schizechinus cf. tuberculatus 




X 








Cyrtechinus verruculatus 




X 




X 




Nudechinus darnleyensis 








X 




Nudechinus multicolor 










X 


Tripneustes gratilla 








X 


X 


Tripneustes preg rat ilia 






X 






Gymnechinus epistichus 








X 


X 


Toxopneustes pileolus 








X 


X 



106 



Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



Species 


Java, Mio- 
Pliocene 


Yule Is, 

Lower 

Pliocene 


Barrow Is, 

Middle 

Miocene 


Australian 

'mainland 1 

species 


Australian 

'reef 

species 


ECHINOMETRIDAE 












Echinometra mathaei 








X 


X 


Heterocentrotus mammillatus 










X 


Echinostrephus aciculatus 










X 


Echinostrephus molaris 










X 














ECHINONEIDAE 












Echinoneus cyclostomus 










X 


Echinoneus abnormalis 










X 














TEMNOPLEURIDAE 












Temnopleurus alexandri 








X 


X 


Temnopleurus toreumaticus 


X 






X 




Salmacis belli 








X 




Salmacis sphaeroides 








X 


X 


Salmacis sphaeroides belli 


X 










Salmacis rarispina 


X 










Salmacis bicolor 


X 










Temnotrema macleayana 


X 


X 








Temnotrema bothryoides 








X 




Temnotrema siamense 








X 


X 


Temnotrema phoenissa 










X 


Opechinus cf. collignoni 


x 










Opechinus cf. cheribonensis 


X 










Opechinus madurae 


X 










Opechinus percultus 


X 










Opechinus percultus var. 
oliaoporus 


X 










Martinechinus molengraaffi 


X 










Microcyphus sp. 


X 










Desmechinus rembangensis* 


X 










Desmechinus erbi 


X 










Mespilia globulus 








X 


X 


Temnopleurid sp. 










X 














ARACHNOIDIDAE 












Arachnoides placenta 








X 
















CLYPEASTERIDAE 












Clypeaster reticulatus 


X 


X 




X 




Clypeaster humilis 


X 


X 




X 




Clypeaster latissimus 




X 




X 




Clypeaster telurus 








X 




Clypeaster blumenthali 


X 










Clypeaster brevipetalus 


X 










Clypeaster butleri 






X 






Clypeaster cf. malumbang- 
ensis 


X 










Clypeaster sp. A 


X 











Proc. Linn. Soc. N.S.W., 125, 2004 



107 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 



Species 


Java, Mio- 
Pliocene 


Yule Is, 

Lower 

Pliocene 


Barrow Is, 

Middle 

Miocene 


Australian 

'mainland' 

species 


Australian 

'reef 

species 


CLYPEASTERIDAE (Cont) 












Clypeaster sp. B 


X 






















LAGANIDAE 












Laganum decagonale 


X 


X 






X 


Laganum depressum 


X 


X 




X 


X 


Laganum depressum var. 
sinaiticum 




X 








Laganum depressum var. 
delicatum 




X 








Laganum herklotzi 


X 










Peronella lesueuri 


X 






X 


x ; 


Peronella orbicularis 








X 


x ! 


Sismondia javana 


X 






















ASTRICLYPEIDAE 












Echinodiscus angulosus 


X 










Echinodiscus tenuissimus 








X 




Echinodiscus sp. 


X 






















FIBULARIIDAE 












Fibularia crispa 


X 










Fibularia cf. scabra 


X 










Fibularia rhedeni 


X 










Fibularia ovulum 










X 


Fibularia volva 








X 




Fibularid sp. 






X 




X 


Echinocyamus cf. cribellum 


X 










Echinocyamus sp. 


X 






















ECHINOLAMPADIDAE 












Echinolampas ovatus 


X 










Echinolampas elevatus 


X 










Echinolampas depressus 


X 










Echinolampas tumulus 






X 


















PLIOLAMPADIDAE 












Pliolampas minutus 


X 










Pliolampas javanus 


X 










Pliolampas elevatus 


X 






















HEMIASTERIDAE 












Hemiaster cf. eupetalum 


X 










Opissaster sp. 


X 






















SPATANGIDAE 












Maretia planulata 


X 


X 




X 


X 


Maretia cordata 




X 








Maretia bandaensis 


X 










Maretia mojsvari 


X 











108 



Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



Species 


Java, Mio- 
Pliocene 


Yule Is, 

Lower 

Pliocene 


Barrow Is, 

Middle 

Miocene 


Australian 

'mainland' 

species 


Australian 

'reef 

species 


PALAEOSTOMATIDAE 












Palaeostoma kairukuensis 




X 




















BRISSIDAE 












Metalia spatagus 








X 


X 


Metalia sternalis 










X 


Brissus latecarinatus 


X 






X 


X 


Brissus sp. 


X 










Rhynobrissus hemiasteroides 








X 




Eupatagus (Eupatagus) 
pulchellus 


X 


X 








Eupatagus affinis 


X 










Eupatagus sp. 


X 






















LOVENIIDAE 












Breynia aff. carinata 






X 






Breynia australasiae 








X 




Breynia paucituberculata 


X 










Breynia sp. a 


X 










Breynia sp. b 


X 










Lovenia elongata 








X 


X 














SCHIZASTERIDAE 












Schizaster (Schizaster) 
lacunosus 








X 


X 


Schizaster (Schizaster) 
alphonsei 




X 








Schizaster (Schizaster) 
compactus 








X 




Schizaster (Schizaster) aff. 
compactus 






X 






Schizaster (Schizaster) sp. A 










X 


Schizaster subrhomboidalis 


X 










Schizaster progoensis 


X 










Schizaster cf. pratti 


X 










Schizaster excavatus 


X 










Schizaster jeanneti 


X 










Schizaster sp. 1 


X 










Schizaster sp. 2 


X 










Schizaster sp. 3. 


X 










Schizaster sp. 


X 










Proraster jukesii 


X 










Moira lethe 








X 




Hemifaorina tuber 


X 











Proc. Linn. Soc. N.S.W., 125, 2004 



109 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 



ERRATA 

The following figures are replacements for: 



Figure 2 (page 128) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule 
Island, Papua New Guinea: Clypeasteroida. Proceedings of the Linnean Society of New South Wales 124, 
125-136. 



Figure 1 (page 155) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule 
Island, Papua New Guinea: Spatangoida. Proceedings of the Linnean Society of New South Wales 124, 153- 
162. 



1 10 Proc. Linn. Soc. N.S.W., 125, 2004 



ID. LINDLEY 




Replacement Figure 2 (page 128) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower 
Pliocene), Yule Island, Papua New Guinea: Clypeasteroida. Proceedings of the Linnean Society of New 
South Wales 124, 125-136. 



Proc. Linn. Soc. N.S.W., 125, 2004 



111 



ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 



1 12 Proc. Linn. Soc. N.S.W., 125, 2004 



ID. LINDLEY 




Replacement Figure 1 (page 155) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower 
Pliocene), Yule Island, Papua New Guinea: Spatangoida. Proceedings of the Linnean Society of New South 
Wales Y2A, 153-162. 



Proc. Linn. Soc. N.S.W., 125, 2004 



113 



114 



Some living and fossil echinoderms from the Bismarck 
Archipelago, Papua New Guinea, and two new echinoid species 

I.D. LlNDLEY 

Department of Geology, Australian National University, Canberra, A.C.T. 0200. 
(lindley @ geology.anu.edu.au) 



Lindley, I.D. (2004). Some living and fossil echinoderms from the Bismarck Archipelago, Papua New 
Guinea, and two new echinoid species. Proceedings of the Linnean Society of New South Wales 125, 
115-139. 

Starfish and sea-urchin records of the Bismarck Archipelago, Papua New Guinea, are scattered throughout 
the literature of the past 160 years. This paper lists the region's valid starfish and sea-urchin species records 
contained in the literature. In addition, records of 17 species of starfish and sea-urchins from material in the 
Department of Geology, Australian National University and the East New Britain Historical and Cultural 
Centre collections are included, with descriptions of two new sea-urchin species, the schizasterid Schizaster 
(Paraster) ovatus sp. nov. and the echinometrid Heliocidaris robertsi sp. nov. Some Tertiary echinoids 
from the region are described for the first time, namely Stereocidaris cf. squamosa Mortensen 1928 (Lower- 
Middle Miocene: Manus Island), Stereocidaris sp. (Pliocene: east New Britain), Phyllacanthus sp. (Pliocene: 
east New Britain) and Echinoneus sp. (Pleistocene-Holocene: Tanga Group, New Ireland). 

Manuscript received 22 August 2003, accepted for publication 22 October 2003. 

KEYWORDS: Asteroidea, Bismarck Archipelago, East Indies, Echinoidea, Extant, Fossil, Papua New 
Guinea, West Pacific. 



INTRODUCTION 

The Bismarck Archipelago, northern Papua 
New Guinea (PNG), encompasses the islands of New 
Britain, Bougainville, New Ireland and adjacent groups 
(Tabar, Lihir, Tanga and Feni), St. Matthias Group, 
the Admiralty Group, including Manus Island, and the 
surrounding waters of the Bismarck Sea (Fig. 1). It 
lies along the easternmost boundary of the East Indian 
Faunal Province. To the east and southeast lies the West 
Pacific Ocean or Melanesia faunal province (Endean 
1957 and A.M. Clark and Rowe 1971, respectively). 

Knowledge of the extant starfishes and sea- 
urchins (Echinodermata: Asteroidea and Echinoidea, 
respectively) from the Bismarck Archipelago 
comprises records scattered throughout a diverse 
literature of the past 160 years. The earliest described 
asteroid is Echinaster eridanella Miiller and Troschel 
1842 (= Echinaster luzonicus Gray 1840) with a type 
locality in New Ireland. Sladen (1889) and A. Agassiz 
(1879 1881) described the asteroids and echinoids, 
respectively, collected during the 1873-76 voyage of 
H.M.S. Challenger. This expedition passed through the 
Admiralty Group and retrieved two new deep-water 
echinoids in the Bismarck Sea (the arbaciid 



Pygmaeocidaris prionigera (A. Agassiz 1879) and the 
temnopleurid Prionoechinus sagittiger A. Agassiz 
1 879), at a site between the Admiralty Group and New 
Guinea. Loriol (1891) described additional asteroids 
from the archipelago, including Nardoa finschi de 
Loriol 1891 (= Nardoa tuberculata Gray 1840) and 
Nardoa mollis de Loriol 1891 (= Nardoa 
novaecaledoniae Perrier 1875), both with type 
localities in New Britain. Bell (1899) described the 
non-holothurian echinoderms collected by Arthur 
Willey during his 1895-97 visit to New Britain and 
the Loyalty Islands (Willey 1902). H.L. Clark (1925) 
redescribed several of Willey' s Bismarck Archipelago 
echinoids and erected two new species (the arbaciid 
Coelopleurus elegans (Bell 1899) and the diadematid 
Micropyga nigra H.L. Clark 1925) with type localities 
in New Britain. H.L. Clark (1946) and A.H. Clark 
(1954) recorded additional asteroids and echinoids 
from the archipelago. Struder (1876, 1880) and H.L. 
Clark (1908) provided descriptions of the extant 
echinoderm fauna of west New Guinea, a region 
contiguous with the Bismarck Archipelago. 

During the past 20 years echinoderm research 
in the region has concentrated on the biology of 
asteroids and comatulid crinoids at Hansa Bay and 
Madang, on the southern shores of the Bismarck Sea 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 




AUSTRALIA 



Figure 1. Locality map showing the Bismarck Archipelago, Papua New Guinea, and other localities 
discussed in text. 



(Britayev et al. 1999; Bouillon and Jangoux 1984; 
Eeckhaut et al. 1996; Messing 1994). 

This paper describes some extant asteroids and 
echinoids from the Bismarck Archipelago. It also 
provides for the first time, a tabulation of previously 
reported asteroid and echinoid occurrences (Tables 1 
and 2 respectively; tables located after the reference 
list) in the region. Several Tertiary echinoids from the 
archipelago are also described. The author is not aware 
of any previous description of the region's fossil 
echinoderm fauna. 

The specimens described in this paper were 
collected between 1981-2003, and do not necessarily 
represent the results of thorough, methodical site 
collections. The Cape Gazelle, east New Britain, 



locality encompasses any one of a number of nearby 
localities, including Tovarur Plantation, Reiven Beach 
and southeastern Tokua Airport. Full systematic 
descriptions are provided for all fossil species while 
in most cases, only brief remarks concerning the 
significance of occurrences are provided for extant 
species. Specimens prefixed ANU are housed in the 
Department of Geology, The Australian National 
University; specimens prefixed B are housed in the 
East New Britain Historical and Cultural Centre, 
Kokopo, East New Britain Province, PNG. 
Terminology and classification used herein follows that 
of the Treatise on Invertebrate Paleontology and A.M. 
Clark and Rowe (1971). 



Proc. Linn. Soc. N.S.W., 125, 2004 



ID. LINDLEY 



SYSTEMATIC DESCRIPTIONS 

Class STELLEROIDEA Lamarck 1816 
Subclass ASTEROIDEA de Blainville 1830 
Order VALVATIDA Perrier 1884 
Suborder GRANULOSINA Perrier 1894 
Family OPHIDIASTERIDAE Verrill 1867 
Genus LINCKIA Nardo 1834 

Synonymy 

Cribella Agassiz 1835 (non Forbes 1841). 
Acalia Gray 1840. 
Catantes, Undina Gistl 1847. 

Type species 

Linkia typus Nardo 1834 (= Asterias laevigatus Linnaeus 1758) by original designation. 

Linckia multifora (Lamarck 1816) 

Synonymy 

Asterias multifora Lamarck 1816, p. 565. 
Linckia leachi Gray 1840, p. 285: Mauritius. 
Linckia costae Russo 1894, p. 163: Daret Is., Red Sea. 

Materials and locality 

Two specimens, ANU 60651-2, collected at Ralum, Blanche Bay, East New Britain Province, PNG. 

Remarks 

Linckia multifora (Lamarck 1816) is widely distributed throughout the Indo-Pacific, from the Red Sea to 
the Hawaiian Islands (A.M. Clark and Rowe 1971). This record is the first from New Guinea. 

Family OREASTERIDAE 

Genus PROTOREASTER Doderlein 1916 

Type species 

Asterias nodosa Linnaeus 1758, p. 420, by subsequent designation. 

Protoreaster nodosus (Linnaeus 1758) 

Synonymy 

Oreaster nodosus, Bell 1884, p. 70; H.L. Clark 1908, p. 280; Fisher 1911, p. 346; H.L. Clark 1921, p. 

31. 
Pentaceros nodosus, Bell 1899, p. 136. 
Protoreaster nodosus, Doderlein 1916, p. 420; H.L. Clark 1946, p. 106; A.M. Clark and Rowe 1971, 

p. 34, 54; Rowe and Gates 1995, p. 106. 

Material and locality 

Single beach worn specimen, ANU 60650, collected at Ralum, Blanche Bay, East New Britain 
Province, PNG. 

Remarks 

Protoreaster nodosus (Linnaeus 1758) is a common East Indian starfish with a range extending to the 
West Pacific (Caroline Islands) (H.L. Clark 1946; A.M. Clark and Rowe 1971). Bell (1899) previously described 



Proc. Linn. Soc. N.S.W., 125, 2004 1 n 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 



the species from the collections of Arthur Willey in Blanche Bay. H.L. Clark (1908, p. 280) described variations 
in specimens from several west New Guinea localities (Humboldt Bay, Sorong, Ansus, Jappen Island). Although 
the present specimen (R/r = 20/9 mm) has lost most of its granules and abactinal plates, it is identified as a 
juvenile P. nodosus (L.M. Marsh, pers. comm.). It is similar to a specimen of P. nodosus in the Western Australian 
Museum (WAM 599-76: R/r = 27/1 1 mm) collected by L.M. Marsh from Pulau Langkai, off south Sulawesi, 
Indonesia. Two juvenile specimens (R = 11-12 mm) from the Andaman Islands, figured and described by 
Koehler (1910: plate XVI, fig. 1) as Anthenea sp., are also very similar to ANU 60650. These specimens may 
also be P. nodosus (L.M. Marsh, pers. comm.). 

Class ECHINOIDEA Leske 1778 

Subclass PERISCHOECHINOIDEA M'Coy 1849 

Order CIDAROIDA Claus 1880 

Family CIDARIDAE Gray 1825 

Subfamily STEREOCIDARrNAE Lambert 1900 

Genus STEREOCIDARIS Pomel 1883 

Synonymy 

Typocidaris Pomel 1883 
Phalacrocidaris Lambert 1902 
Anomocidaris Agassiz and Clark 1907 

Type species 

Cidaris cretosa Mantell 1835; subsequent designation Lambert and Thiery 1909 (Feb., p. 31; non 
Mar., where C. merceyi was designated, p. 152). 

Stereocidaris cf. squamosa Mortensen 1928 
Figs 2, 3a 

Synonymy 

Stereocidaris indica Bell 1909, p. 21; H.L. Clark 1925, p. 26. 
Stereocidaris squamosa Mortensen 1928b, p. 70; Mortensen 1928a, p. 245. 





Figure 2. Stereocidaris cf. squamosa Mortensen 1928. Lower-Middle 
Miocene, Manus Island, Manus Province. 2a-b, plating diagrams at 
ambitus for ambulacrum, interambulacrum. Abbreviations: btr basal 
terrace; it inner tubercle; m mamelon; mt marginal tubercle; r ridge; 
scr scrobicular ring; w wall. 



Material 

An incomplete specimen 
ANU 60638, with an 
interambulacral plate and portion 
of ambulacral series. 

Locality and horizon 

Village of Drankei, west 
bank of Wari River, central 
southern Manus Island, Manus 
Province, PNG. Grid reference 
060612 Lorengau 1: 100 000 Sheet 
8393 (Edition 1). The collection 
horizon is an outlier of the (lower) 
Mundrau Limestone. A sample of 
limestone from a nearby outlier of 
the Mundrau Limestone at 
Metawarei village, 0.5 km 
northwest of Drankei village, 
contained a foraminiferal 
assemblage of mid or upper Tfl 
age, and suggests a late Lower 



118 



Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 




Figure 3. Tertiary echinoids from the Bismarck Archipelago. Stereocidaris cf. squamosa Mortensen 1928. 
Lower-Middle Miocene, Manus Island, Manus Province. 3a, ANU 60638, incomplete interambulacral 
plate with large tubercle and part of adjacent ambulacral plating (refer to Figs 2a, b for plating diagram). 
Bar scale = 2.5 mm. Stereocidaris sp. Pliocene, Sikut River area, East New Britain Province. 3b, ANU 
60639, primary spine. Bar scale = 2.5 mm. Phyllacanthus sp. Pliocene, Mevelo River area, East New 
Britain Province. 3c, ANU 60637, proximal portion of primary spine. Bar scale = 5 mm. Echinoneus sp. 
Pleistocene-Holocene, Boang Island, Tanga Group, New Ireland Province. 3d, ANU 60640, aboral view 
of worn specimen. Bar scale = 5 mm. 



Miocene or earliest Middle Miocene age for the unit (Francis 1985). 

Description 

Test size and shape unknown. 

Ambulacra sinuate, rather broad, ca. 39% of width of interambulacra. Interporiferous zone about twice 
width of a pore-zone. Interporiferous zone with distinctly vertical series of marginal tubercles and inner tubercles; 
marginal tubercles slightly larger than those of inner series. Pores are rather small, circular, nonconjugate and 
separated by a broad wall; ridge low and narrow. The arrangement of pores and tubercles in ANU 60638 is 
strikingly similar to that described for Stereocidaris squamosa Mortensen 1928 by Mortensen (1928a: 245 and 
Plate LXX, fig. 7). 

Interambulacral plate higher than broad (height:width = 8.0:5.5) with aureole of moderate size but not 
very deep, well separated. The very high plate of ANU 60638 exhibits a broad miliary covered space between 



Proc. Linn. Soc. N.S.W., 125, 2004 



119 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 

successive aureoles, one unequal to the other, indicating it to be from an upper interambulacral, or aboral, 
position on test (Fig. 2). Mamelon apparently large, the part of plate is damaged in ANU 60638. Edge of 
aureole is not raised, and the scrobicular tubercles are not prominent. Outside the scrobicular ring, the 
interambulacral plate has a sparse to moderate covering of tubercles of similar size to scrobicular tubercles. On 
the adradial edge of plate there are a few secondary tubercles outside the scrobicular ring. 

Apical and periproctal systems unknown. 

Details of primary and secondary spines unknown. 

Remarks 

Stereocidaris Pomel 1883 is a well characterised genus with both fossil and extant species (Mortensen 
1928a; Chapman and Cudmore 1934). The genus is distinctive for its usually very high interambulacral plates, 
ambulacra that are generally conspicuously sinuate and nonconjugate pores (Mortensen 1928a; Fell 1966). The 
oldest occurrence of the genus is from the Cretaceous of Europe. In the Tertiary it is known from the Eocene of 
Europe and Australia, Oligocene of New Zealand, Miocene of Australia and Indonesia, and Pliocene of Australia 
and New Zealand (Mortensen 1928a; Chapman and Cudmore 1934; Fell 1966). Mortensen (1928a) noted the 
lack of fossil Stereocidaris from the Indo-Pacific, with K. Martin's (1918) record of the occurrence of a spine 
of Dorocidaris papillata (= Stereocidaris) from the Miocene of Java the only known fossil. This may well 
represent a collection bias. Extant species of Stereocidaris, numbering 15, with nine subspecies, are distributed 
throughout the Indo-Pacific, including southeast Africa (Mortensen 1928a). Notably the genus has not been 
recorded from Australasian seas (HL. Clark 1925; Fell 1966). 

The Manus Island specimen, represented by a small fragment of plating from an adapical position, is 
tentatively assigned to Stereocidaris squamosa Mortensen 1928. As already noted, the striking resemblance of 
the ambulacral and interambulacral plating of this single specimen to that described by Mortensen (1928a) for 
S. squamosa cannot be ignored. Stereocidaris squamosa is an extant species recorded from 270 m depth on the 
Saya de Malha Bank (10° 30'S), about 800 km southeast of the Seychelles in the Indian Ocean (Mortensen 
1928a). The species has a small-moderate sized test that ranges in diameter from 30-47 mm, with height from 
18.5-29 mm (Mortensen 1928a). Longest spines range in length from ca. 50-59 mm. The late Lower Miocene/ 
earliest Middle Miocene Manus Island occurrence represents the first fossil occurrence of test remains of 
Stereocidaris from the Indo-Pacific. 

Stereocidaris sp. 

Fig. 3b 

Material 

An isolated fragmentary spine, ANU 60639. 

Locality and horizon 

Collected '5 km east of the intake structure of the Warangoi hydro-scheme' (Lindley unpubl. field 
notes), in the headwaters of Matuli Creek, a tributary of the Warangoi River, Sikut area, northeastern Gazelle 
Peninsula, East New Britain Province, PNG. Grid reference 081961 Merai 1: 100 000 Sheet 9388 (Edition 1). 
The collection horizon is from the Sinewit Formation, of Mio-Pliocene age (Lindley 1988). However, fossil 
evidence and a K-Ar radiometric age from the Sikut and adjacent areas, indicates the formation in this area is 
restricted to the Pliocene (Read 1965; B. McGowran in Lindley 1988; Lindley 1988; Corbett et al. 1991). 

Description 

No test fragments which belong to this species haye been identified. 

Primary spine cylindrical, distinctly fusiform, tapering, point not widened. Spine length 15 mm, with 
maximum diameter of 2 mm occurring about 1/2 distance from proximal end. The shaft with about 16 series of 
low rounded warts; only towards the point do they assume the shape of low rounded ridges. The collar is only 
0.75-1 mm long, slightly increasing in thickness towards inconspicuous milled ring. Neck is equal in length to 
collar. 

Remarks 

The primary spines of cidaroids possess a distinctive structure with a compact outer or cortex layer 



120 Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



covering all except the collar and enveloping a central core consisting of an irregular calcareous meshwork 
(Mortensen 1928a). The cortex layer is found only in a few other echinoids, mainly the salenids, and spinule 
and wart ornament along the shaft is formed by this alone (Mortensen 1928a). Mortensen (1928a: 50) considered 
that primary spine shape and structure is of considerable use in cidaroid classification, both at specific and 
generic levels. 

The primary spine ANU 60639 is identified as that of a cidaroid by its spine shape and its possession of 
an outer cortex layering. The nature of the inner central core meshwork is clearly visible on the spine collar. The 
nature of wart development, the number of longitudinal series, and their distal transition to low rounded ridges, 
bears a strong resemblance to that seen in the primary spines of some extant species of Stereocidaris, including 
Stereocidaris grandis (Doderlein) and Stereocidaris hawaiiensis Mortensen 1928b, found only in Japanese 
seas and Hawaiian seas, respectively (cf. Mortensen 1928a: Plate XIX, fig. 5 and XXI, fig. 5, respectively). 

Subfamily RHABDOCIDARINAE Lambert 1900, emended Fell 1966 
Genus PHYLLACANTHUS Brandt 1835 

Synonymy 

Leiocidaris Desor 1885, p. 48. 

Type species 

Cidarites (Phyllacanthus) dubia Brandt 1835, p. 67, by original designation. 

Phyllacanthus sp. 

Fig. 3c 

Material 

One isolated fragmentary spine, ANU 60637. 

Locality and horizon 

Collected in stream float from an unnamed large western tributary of Mevelo River, Lakit Range, 
southwestern Gazelle Peninsula, East New Britain Province, PNG. Grid reference 660623 Pondo 1:100 000 
Sheet 9288 (Edition 1). Lakit Limestone, Pliocene (Lindley 1988). 

Description 

No test fragments which belong to this species have been identified. 

Proximal portion of primary spine moderately thick, cylindrical, fusiform, with a maximum diameter of 
8.0 mm. Details of distal shaft unknown. Details of spine base, milled ring and collar unknown. Spine swells 
rapidly above the collar. Surface of shaft is finely and uniformly granulated (not visible to the naked eye), the 
granules forming numerous (> 50) longitudinal series along length of spine. 

Remarks 

Lindley (2003b) described the spines of Phyllacanthus imperialis var. javana K. Martin 1885 and 
Phyllacanthus sp. from the Lower Pliocene Kairuku Formation, Yule Island. Unfortunately, the characters 
diagnostic of these species, including spine collar length and the number of ridges on the distal part of the spine, 
are not visible on ANU 60637. 

Subclass EUECHINOIDEA Bronn 1860 
Superorder ECHINACEA Claus 1876 
Order TEMNOPLEUROIDA Mortensen 1942 
Family TOXOPNEUSTIDAE Troschel 1872 
Genus TOXOPNEUSTES A. Agassiz 1841 

Synonymy 

Boletia Desor 1846, p. 362. 



Proc. Linn. Soc. N.S.W., 125, 2004 121 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 

Type species 

Echinus pileolus Lamarck 1816, p. 45, by original designation. 

Toxopneustes pileolus (Lamarck 1816) 

Synonymy 

Echinus pileolus Lamarck 1816, p. 45. 

Toxopneustes pileolus, A. Agassiz 1841, p. 7; HI. Clark 1925, p. 123; Mortensen 1943a, p. 472; 

A.M. Clark and Rowe 1971, p. 156; Rowe and Gates 1995, p. 258. 
Mortensen (1943a: 472) lists additional synonymies. 

Material and locality 

Single naked test, B20022, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 

Remarks 

Toxopneustes pileolus (Lamarck 1816) is widely distributed throughout the Indo-West Pacific 
(Mortensen 1943a; A.M. Clark and Rowe 1971; Miskelly 2002). 

Genus TPJPNEUSTES L. Agassiz 1841 

Type species 

Echinus granulans Lamarck 1816, p. 44, by original designation. 

Tripneustes gratilla (Linnaeus 1758) 

Synonymy 

Echinus gratilla Linnaeus 1758, p. 664. 

Tripneustes gratilla, HL. Clark 1925, p. 124; Mortensen 1943a, p. 500; A.M. Clark and Rowe 1971, 

p. 156; Rowe and Gates 1995, p. 259. 
Mortensen (1943a: 500) lists additional synonymies. 

Material and locality 

Single naked test, B20023, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 

Remarks 

Tripneustes gratilla (Linnaeus 1758) is widely distributed throughout the Indo-West Pacific (Mortensen 
1943a; A.M. Clark and Rowe 1971). Previous records from the Pacific include the Marshall Islands, Norfolk 
Island, Hawaiian Islands, Kermadec Islands, Solomon Islands, Fiji and Hood Lagoon, south coast of Papua 
(HL. Clark 1925; Mortensen 1943a; A.M. Clark and Rowe 1971; Miskelly 2002). 

Order ECHTNOIDA Claus 1876 
Family ECfflNOMETRIDAE Gray 1825 
Genus ECHINOMETRA Gray 1825 

Synonymy 

Ellipsechinus Liitken 1864, p. 165. 
Plagiechinus Pomel 1883, p. 78. 
Mortensenia Doderlein 1906, p. 233. 

Type species 

Echinus lucunter Linnaeus 1758, p. 665, by original designation. 



122 Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 

Echinometra mathaei (de Blainville 1825) 

Synonymy 

Echinus lucunter Lamarck 1816, p. 50 (non E. lucunter Linnaeus). 

Echinometra mathaei, H.L. Clark 1925, p. 143; H.L. Clark 1932, p. 216; Mortensen 1943b, p. 381; 

H.L. Clark 1946, p. 332; A.M. Clark and Rowe 1971, p. 157; Rowe and Gates 1995, p. 211. 
Mortensen (1943b: 381) lists additional synonymies. 

Material and localities 

Fourteen naked tests from Gargaris village, northern coast of Malendok Island, Tanga Group, New 
Ireland Province, PNG; one partly naked test from beach at Ralum, Blanche Bay, East New Britain Province, 
PNG; one naked test from Penlolo village, south coast of New Britain, West New Britain Province, PNG; one 
naked test, B 20016, from Cape Gazelle, New Britain, East New Britain Province, PNG. 

Remarks 

Echinometra mathaei (de Blainville 1825) is a long ranging species, recorded from late Lower Miocene- 
early Middle Miocene rocks in the western and eastern Mediterranean Sea (Negretti et al. 1990). Extant E. 
mathaei is one of the most widely distributed echinoids, occurring throughout tropical-subtropical waters of the 
Indo-West Pacific (Mortensen 1943b; A.M. Clark and Rowe 1971). H.L. Clark (1908) recorded the species 
from Sorong, west New Guinea and Miskelly (2002) recorded it from the Solomon Islands. This record indicates 
a wide distribution throughout the Bismarck Archipelago (Tanga Group, New Ireland; Blanche Bay, New 
Britain; and south coast New Britain). 

Genus HETEROCENTROTUS Brandt 1835 

Synonymy 

Acroladia L. Agassiz and Desor 1846, p. 373. 

Type species 

Echinus mamillatus Linnaeus 1758, p. 664, by subsequent designation of Pomel 1883, p. 77. 

Heterocentrotus mammillatus (Linnaeus 1758) 

Synonymy 

Echinus mamillatus Linnaeus 1758, p. 664. 

Heterocentrotus mammillatus, H.L. Clark 1925, p. 147; Mortensen 1943b, p. 409; H.L. Clark 1946, p. 

333; A.M. Clark and Rowe 1971, p. 158; Rowe and Gates 1995, p. 213. 
Mortensen (1943b: 409) lists additional synonymies. 



Material and locality 

A single naked test, B 20017, and unlabelled isolated spines (housed in the East New Britain Historical 
and Cultural Centre, Kokopo) from Cape Gazelle, New Britain, East New Britain Province, PNG; an isolated 
primary spine, ANU 60648, from Nosnos village, Boang Island, Tanga Group, New Ireland Provine, PNG. 

Remarks 

Heterocentrotus mammillatus (Linnaeus 1758) is widely distributed throughout the Indo-Pacific, from 
the Gulf of Suez and Madagascar to the Hawaiian Islands and Fiji (Mortensen 1943b). It is recorded from the 
Solomon Islands by Miskelly (2002). The largest test of H. mammillatus noted by Mortensen (1943b) has a 
long diameter of 82 mm, with most individuals having diameters of 72 mm or less. The long diameter of the 
Cape Gazelle test is 72 mm. The Tanga spine has a length of 74 mm and, given that the primary spines of H. 
mammillatus usually do not exceed the long diameter of the test (Mortensen 1943b), appears to have come from 
a relatively large individual. 



Proc. Linn. Soc. N.S.W., 125, 2004 123 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 



Genus HELIOCIDARIS L. Agassiz and Desor 1846. 

Synonymy 

Toxocidaris A. Agassiz 1863, p. 22. 

Type species 

Echinus tuberculatus Lamarck 1816, p. 50, by original designation. 



Diagnosis 

Low hemispherical echinoids, widest at circular ambitus. Ambulacral plates with 7 or more pore-pairs to 
each plate; arcs may be irregularly double; expanded poriferous tracts of the flattened adoral surface are petaloid. 
Oculars I and IV usually insert. Gill-slits are shallow (Philip 1965; Fell and Pawson 1966). 

Remarks 

Heliocidaris L. Agassiz and Desor 1846 is distributed along the southern coasts of Australia, northern 
New Zealand, Kermadec Islands and Lord Howe Island (Mortensen 1943 a). Two species are included in the 
genus by Mortensen (1943a), viz: Heliocidaris tuberculata (Lamarck 1816) and Heliocidaris ery thro gramma 
(Valenciennes 1846) and, given their similar morphologies, he has questioned whether they are really 
conspecific. Anthocidaris Liitken 1864 is a closely allied genus (only known species Anthocidaris 
crassispina [A. Agassiz 1863]) from the coasts of southern Japan and China, distinguished from Heliocidaris 
by the spicules of the tubefeet (Mortensen 1943a). On the status of Anthocidaris, Mortensen (1943a: 328) 
questioned whether the genus should be merged into Heliocidaris. Philip (1965) described the only known 
fossil representative of the genus, Heliocidaris ludbrookae Philip 1965 from the Lower-early Middle 
Miocene (Longfordian-Batesfordian) of southeastern Australia. 

Heliocidaris robertsi sp. nov. 
Figs 4, 5a-e 

Diagnosis 

Test low hemispherical, somewhat inflated above. Ambulacral plates with 12 pore-pairs per plate; ambital 
and aboral pore-arcs doubled. Ambulacral and interambulacral plates relatively large; each bearing a primary 
tubercle and numerous secondary tubercles; aureoles of primaries not in contact. Primary tubercles of ambital 
and aboral ambulacral plates with an aborally positioned secondary tubercle. 

Etymology 

Named for Mr Michael Roberts, amateur conchologist of Kokopo, East New Britain Province, PNG. 

Material and locality 

Single naked test, ANU 
60654, from the vicinity of 
Cape Gazelle, New Britain, 
East New Britain Province, 
PNG. 



Description 

Test low hemispherical, 
somewhat inflated above, 
widest at circular ambitus. The 
oral side is flattened, scarcely 
sunken towards the peristome. 
Only specimen of 38 mm 
diameter. 

The pore zones are 
conspicuously petaloid on the 





Figure 4. Heliocidaris robertsi sp. nov. Cape Gazelle area, East New Britain 
Province. 4a-b, plating diagrams at ambitus for interambulacrum, 
ambulacrum. 



124 



Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 




Figure 5. Heliocidaris robertsi sp. nov. Cape Gazelle area, East New Britain Province. 5a-e, ANU 60654, 
aboral, oral, lateral views. Bar scale = 10 mm; ambulacral plating at ambitus (refer to Fig. 4b for plating 
diagram). Bar scale = 5 mm; apical disc. Bar scale = 2.5 mm. 



Proc. Linn. Soc. N.S.W., 125, 2004 



125 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 

oral surface, about 1 .5-2 times the width of interporiferous zone. The pore-series in this area are almost horizontal 
and are separated by secondary tubercles forming a single prominent vertical series; scattered miliary tubercles 
are also present. In the ambital region there are 12 pore-pairs arranged in double arcs (Fig. 4). Above the 
ambitus the pore-zones become much narrower. Primary tubercles in the ambital zone are large, almost as large 
as the interambulacral primaries; aureoles of adjacent primaries in each vertical series widely separated. Sutures 
between adjacent plates are seen very distinctly on the outer adoral side of the boss. Each ambulacral plate at 
and above the ambitus has a prominent secondary tubercle positioned aborally to the primary tubercle; 4-5 
other secondary tubercles are also present. Miliaries tend to be arranged along the perradial sutures of ambital 
and superoambital ambulacrals; elsewhere on each plate only a sparse covering of miliaries is present. 

The interambulacral primaries are large, forming prominent series aborally; their aureoles are distinctly 
separated, leaving a broad space at the upper edge of each plate, occupied by several small tubercles and 
miliaries. Usually sutures between adjacent plates are close to, but do not cross, aureole of successive tubercle. 
In the median space there is on the oral surface and in the ambital region a conspicuous double series of 
secondary tubercles about half the size of the ambulacral primaries. Below the ambitus all the tubercles decrease 
rapidly in size, with the secondaries disappearing, and only the primaries continuing to the peristome. 

The apical system is small, only about 18 percent of the test diameter. There is typically one large 
tubercle on each genital plate, except the very large madreporite, and a scattering of small tubercles over the 
remainder (Fig. 5e). Ocular I and IV are broadly insert. The peristome is very small, about 29 percent of test 
diameter. Gill-slits shallow. 

Details of spines and pedicellarie unknown. 

Remarks 

Heliocidaris robertsi sp. nov. is readily distinguished from H. tuberculata and H. erythrogramma and 
the closely allied A. crassispina by its possession of double pore-arcs on the adoral surface. The double pore- 
arcs of H. robertsi are very similar to those of Heterocentrotus trigonarius (Lamarck 1816), figured by Mortensen 
(1943a: fig. 132c) and Fell and Pawson (1966: fig. 324, 7c). However, any resemblance between the new 
species and H. trigonarius is easily discounted because of the latter' s possession of a distinctly elongated test 
and a significantly larger peristome (51 percent of test diameter). 

The biogeographical position of H. robertsi is noteworthy in that it is the tropical representative of two 
closely allied temperate water genera, Heliocidaris, a very common form restricted to southern Australia and 
New Zealand, and Anthocidaris, an equally common form restricted to Japan and China. 

Pore-arc doubling is almost as strongly developed in other echinometrids including Colobocentrotus 
Brandt 1835 and Zenocentrotus A.H. Clark 193 1, and incipient development may also been seen in Echinometra 
Gray 1825 (Mortensen 1943a: 281). All three genera possess an elliptical or oblong ambitus. The functional 
significance of doubling of pore-arcs in compound plates relates to (a) increasing the area over which tube-feet 
are spread, and thereby increasing respiratory and feeding efficency (Mortensen 1943a; Woods 1958; Durham 
1966; A.M. Clark 1968) and (b) strengthening of the test (Durham 1966). The doubling of pore-arcs on the 
aboral surface of H. robertsi greatly increases the number of tube-feet in this area, not only aiding in improved 
respiration, but allowing it to catch food particles falling onto its upper surface. With such adaptations to its 
upper surface, the echinoid may have been a reef rock borer, inhabiting a hole perhaps several centimetres deep. 

Superorder GNATHOSTOMATA Zittel 1879 
Order HOLECTYPOIDA Duncan 1889 
Suborder ECHINONEINA H.L. Clark 1925 
Family ECHINONEIDAE Agassiz and Desor 1847 
Genus ECHINONEUS Leske 1778 

Synonymy 

Echinanaus Gray 1825, p. 7 (nom. van.). 
Pseudohaimea Pomel 1885, p. 118. 
Koehleraster Lambert and Thiery 1921, p. 331. 

Type species 

Echinoneus cyclostomus Leske 1778, by subsequent designation of H.L. Clark 1917, p. 101. 



126 Proc. Linn. Soc. N.S.W., 125, 2004 



ID. LINDLEY 



Remarks 

Echinoneus Leske 1778 is an Oligocene-Recent form, with some ten fossil species described from the 
Oligocene and Miocene of Europe (Mortensen 1948a; Wagner and Durham 1966). Two Recent species are 
known, viz. Echinoneus cyclostomus Leske 1778 and Echinoneus abnormalis de Loriol 1883, distinguished by 
the presence or absence of imperforate primary tubercles and well developed glassy tubercles. Recent forms are 
distributed throughout the West Indies, Indo-Pacific and Australia. Mortensen (1948a) considered that many of 
the fossil species are very difficult to distinguish and may in fact be Recent E. cyclostomus. 

Echinoneus sp. 

Fig. 3d 

Material 

One poorly preserved test, ANU 60640. 

Locality and horizon 

Nosnos village, Boang Island, Tanga Group, New Ireland Province, PNG. Grid reference 296246 Tanga 
1:100 000 Sheet 9591 (Edition 1). Unnamed poorly compacted bioclastic limestone, Pleistocene-Holocene 
(Wallace et al. 1983). 

Description 

Test ovoid, moderate size, measuring 23 x 17 x 11.5 mm; oral surface weakly concave. Ambulacra 
narrow, not petaloid. Other details of ambulacra unknown. Details of interambulacra unknown. Details of 
tubercles unknown. Apical and periproctal systems unknown. 

Remarks 

The lack of well preserved tubercles on this specimen makes it difficult to assign a species. 

Echinoneus cyclostomus Leske 1778 

Synonymy 

Echinoneus cyclostomus Leske 1778, p. 173; H.L. Clark 1925, p. 177; H.L. Clark 1946, p. 353; 

Mortensen 1948a, p. 75; A.M. Clark and Rowe 1971, p. 158; Rowe and Gates 1995, p. 215. 
Mortensen (1948a: 75) lists additional synonymies. 

Material and locality 

Twelve naked tests, including ANU 60641, from Gargaris village, northern coast of Malendok Island, 
Tanga Group, New Ireland Province, PNG; one naked test, B 20021, from Cape Gazelle, New Britain, East 
New Britain Province, PNG. 

Remarks 

Echinoneus cyclostomus Leske 1778 is the only known case of a (tropical) cosmopolitan echinoid, having 
been recorded from the West Indies, Ascension (but not the African west coast) and the Indo-Pacific-East 
Africa (Zanzibar, Natal), Madagascar to the Pacific islands (Funafuti, Palmyra, Hawaiian Islands), and from 
Japan to Queensland (Great Barrier Reef) and Lord Howe Island (Mortensen 1948a). Miskelly's (2002) record 
of E. cyclostomus from the Solomon Islands represents the nearest previous record to that from the Tanga 
Group and Cape Gazelle. 

Echinoneus abnormalis de Loriol 1883 

Synonymy 

Echinoneus abnormalis de Loriol 1883, p. 41; H.L. Clark 1917, p. 102; H.L. Clark 1925, p. 176; 

Mortensen 1948a, p. 80; A.M. Clark and Rowe 1971, p. 158. 
Koehleraster abnormalis Lambert and Thiery 1921, p. 331. 



Proc. Linn. Soc. N.S.W., 125, 2004 127 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 



Material and locality 

One naked test, ANU 60641, from Gargaris village, northern coast of Malendok Island, Tanga Group, 
New Ireland Province, PNG. 

Remarks 

This species is represented by a single naked test measuring 30 x 22.5 x 15 mm. Echinoneus abnormalis 
de Loriol 1883 is distinguished from E. cyclostomus by possessing perforated, non-glassy spine tubercles. The 
apical system of the Tanga specimen is distinctly anterior to that of co-occurring specimens of the much more 
common E. cyclostomus. Echinoneus abnormalis has a restricted distribution, known from Mauritius (type 
locality), Kei Islands, Palmyra Island, Banda, Ellice Islands and the Hawaiian Islands (Mortensen 1948a; A.M. 
Clark and Rowe 1971). The recent record of E. abnormalis from the vicinity of Raine Island on the northern 
Great Barrier Reef (Gibbs et al. 1976) represents the first from Australasian waters. The record from the Tanga 
Group is the second from the East Indies. The species is observed to be sympatric with the much more common 
E. cyclostomus in many localities, a fact Gibbs et al. (1976) suggested may have resulted in it having gone 
unrecognised in samples. Mortensen (1948a: 81) considered that the two species probably didn't live together 
at the same localities. Of the 15 specimens of Echinoneus collected from the Malendok Island locality, only one 
was an E. abnormalis, suggesting that in this case, the species' apparent rarity may be related to different niches 
within the same locality. 

Order CLYPEASTEROIDA A. Agassiz 1872 
Suborder CLYPEASTERINA A. Agassiz 1872 
Family CLYPEASTERIDAE L. Agassiz 1835 
Genus CLYPEASTER Lamarck 1801 

Type species 

Clypeaster rosaceus (Linnaeus 1758), by subsequent designation of Desmoulins 1835. 

Clypeaster reticulatus (Linnaeus 1758) 

Synonymy 

Lindley (2003a) lists previous synonymies. 

Material 

Single naked test, B20020, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 

Remarks 

Clarification of Lindley's (2003a) statement on the distribution of Clypeaster reticulatus (Linnaeus 1758) 
is needed. The species is a very common Indo-West Pacific echinoid, distributed in the western Indian Ocean 
and the Red Sea, throughout the East Indies and east into the Pacific Ocean to the Hawaiian Islands (A.M. Clark 
and Rowe 1971). Previous south Pacific records of the species have been made by A. Agassiz (1863), Mortensen 
(1948b) and A.H. Clark (1954) from the Gilbert Islands, New Caledonia and Marshall Islands, respectively. 
Mortensen's (1948b) New Caledonian record has not been confirmed by De Ridder (1986: 29). McNamara and 
Kendrick (1994) have also recorded the species from Barrow Island, northwestern Australia. The species is 
known from fossil in Java (Lower Miocene), Yule Island, PNG (Lower Pliocene), East Africa (Pliocene- 
Pleistocene) and the New Hebrides (Pleistocene) (Mortensen 1948b; Lindley 2003a). 

Family ARACHNOID AE Duncan 1889 
Subfamily ARACHNOIDINAE Duncan 1889 
Genus ARACHNOIDES Leske 1778 

Synonymy 

Echinarchinus Leske 1778, p. 217. 



128 Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



Type species 

Echinus placenta Linnaeus 1758, p. 666, ICZN 1954. 

Arachnoides placenta (Linnaeus 1758) 

Synonymy 

Echinus placenta Linnaeus 1758, p. 666. 

Arachnoides placenta (Linnaeus 1758): L. Agassiz 1841, p. 94; Bell 1899, p. 136; H.L. Clark 1925, p. 

154; H.L. Clark 1946, p. 340; A.M. Clark and Rowe 1971, p. 161; Rowe and Gates 1995, p. 

176. 
Mortensen (1948b) lists additional synonymies. 

Material and locality 

Single naked test, B20018, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 

Remarks 

Arachnoides placenta (Linnaeus 1758) is a common littoral species throughout the East Indies and the 
south Pacific (Mortensen 1948b; A.M. Clark and Rowe 1971). The first record of the species from the Bismarck 
Archipelago is that of Bell (1899) from an unspecified locality in New Britain. 

Suborder LAGANINA Mortensen 1948 
Family LAGANTDAE A. Agassiz 1873 
Genus LAGANUM Link 1807 

Synonymy 

Lagana Gray 1825, p. 427. 

Type species 

Laganum petalodes (= Echinodiscus laganum Leske 1778, p. 204), by original designation. 

Laganum laganum (Leske 1778) 

Synonymy 

Laganum Bonani Klein 1734, p. 25. 
Echinodiscus laganum Leske 1778, p. 204. 
Laganum laganum, Mortensen 1948b, p. 312. 
Laganum depressum, Lindley 2001, p. 130. 
Mortensen (1948b: 312) list previous synonymies. 

Material and locality 

Single test, ANU 60649, from Penlolo village, south coast of New Britain, West New Britain 
Province, PNG. 

Remarks 

Laganum laganum (Leske 1778) is distinct with its pentagonal test with thick, swollen edges, and an 
oblong-elongate periproct situated midway between the mouth and test edge. The species is common in the East 
Indies, and is also recorded from Port Jackson and Tasmania (Mortensen 1948b). Mortensen (1948b) also 
recorded it from the Bismarck Archipelago (Table 2). H.L. Clark (1908) recorded the species from Saonek, 
Waigiou Island, in west New Guinea (Fig. 1) 

Suborder SCUTELLINA Haeckel 1896 
Family ASTRICLYPEIDAE Stefanini 1911 



Proc. Linn. Soc. N.S.W., 125, 2004 129 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 

Genus ECHINODISCUS Leske 1778 

Type species 

Echinodiscus bisperforatus Leske 1778, p. 196. 

Echinodiscus tenuissimus (L. Agassiz in Agassiz and Desor 1847) 

Synonymy 

Lobophora tenuissima L. Agassiz and Desor 1847, p. 136. 

Echinodiscus tenuissimus, Gray 1855, p. 20; H.L. Clark 1914, p. 71; H.L. Clark 1925, p. 171; 

Mortensen 1948b, p. 411; A.M. Clark and Rowe 1971, p. 144 162; Rowe and Gates 1995, p. 

185. 
Mortensen (1948b: 411) lists additional synonymies. 

Material and locality 

Two tests, B 20024 (naked) and B 20025 (with spines), from the vicinity of Cape Gazelle, New 
Britain, East New Britain Province, PNG. 

Remarks 

Echinodiscus tenuissimus (L. Agassiz in Agassiz and Desor 1847) is a widely distributed Indo-West 
Pacific form, occurring throughout the East Indies, northern Australia, southern Japan and the south Pacific 
(Mortensen 1948b; A.M. Clark and Rowe 1971). In the south Pacific, the species is recorded from Tanna, 
Vanuatu, (H.L. Clark 1925) and from New Caledonia (A.M. Clark and Rowe 1971). However, De Ridder 
(1986) only noted the occurrence of Echinodiscus bisperforatus Leske 1778 from New Caledonia. H.L. Clark 
(1925) observed that New Caledonian specimens of E. tenuissimus in the British Museum (Natural History) 
have a form more like E. bisperforatus. The Cape Gazelle specimens have very short lunules, about one quarter 
the length of the radius taken through them, and there is no difference in the tuberculation and spines of the 
ambulacral and interambulacral areas of the oral surface, both diagnostic characters of E. tenuissimus (Mortensen 
1948b; A.M. Clark and Rowe 1971). 

Superorder ATELOSTOMATA Zittel 1879 
Order SPATANGOIDA Claus 1876 
Suborder HEMIASTERTNA Fischer 1966 
Family SCHIZASTERTDAE Lambert 1906 
Genus SCHIZASTER L. Agassiz 1836 

Type species 

Schizaster studeri L. Agassiz 1836, p. 185, by subsequent designation ICZN 1948. 

Remarks 

McNamara and Philip (1980a, b) questioned the familial classification of the spatangoids used by 
Mortensen (1951) and Fischer (1966) and, in particular, the Family Schizasteridae. Within the Schizasteridae 
McNamara and Philip recognized genera sharing the gross morphological test features of Schizaster, viz. a 
posteriorly located apical system, with the apex of the test posterior to this; a long, typically sunken, poriferous 
frontal ambulacrum; and sunken petals, of which the posterior pair are markedly shorter than the anterior ones. 
Within this group, McNamara and Philip (1980a, b) included the genus Schizaster L. Agassiz 1836 (with its 
subgenera Dipneutes Arnaud 1891; Paraster Pomel 1869 and Ova Gray 1825 [= Diploraster Mortensen 1951]); 
Brisaster Gray 1855; Kina Henderson 1975; MoiraL. Agassiz 1 872 (=MoiropsisL. Agassiz 1881); and Proraster 
Lambert 1895 (= Hypselaster Clark 1917). The author accepts their emended diagnosis for Schizaster. 

Subgenus PARASTER Pomel 1869 

Type species 

Schizaster gibberulus L. Agassiz 1847, by original designation of Pomel 1869, p. 14. 



130 Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



Diagnosis 

Species of Schizaster with a small to moderate sized test, with a shallow frontal sinus. Apical system 
slightly posterior of centre. Frontal ambulacrum shallow with pore pairs inclined at about 45° and arranged in 
single rows. Anterior petals almost straight, diverging at an angle up to 110° (McNamara and Philip 1980a). 

Remarks 

There is difficulty in placing the Cape Gazelle species firmly within McNamara and Philip's (1980a) 
subgenus Paraster Pomel 1869. This is particularly in relation to details of the anterior petals, their flexed 
nature and 80° angle of divergence, both characters diagnostic of subgenus Schizaster L. Agassiz 1836. The 
frontal ambulacrum does not possess the steeper sided walls typical of species referred to Schizaster (Schizaster) 
(McNamara and Philip 1980a). Furthermore, McNamara and Philip (1980a) noted that species referred to 
Schizaster (Schizaster) possess a more elongate, narrower test than those assigned to Paraster. The Cape Gazelle 
species is assigned to Schizaster (Paraster) by its possession of a small test, shallow frontal sinus, apical system 
slightly posterior of centre and shallow frontal ambulacrum with pore pairs inclined at about 45°. The species 
is probably morphologically transitional between the Paraster and Schizaster morphotypes. 

Schizaster (Paraster) ovatus sp. nov. 
Figs 6a-d 



Diagnosis 

A small species of Schizaster (Paraster) with a moderately depressed, ovoid test; apical system is 55 
percent of test length from anterior, with four genital pores. Anterior ambulacrum relatively narrow and shallow; 
pore pairs inclined at about 45° and arranged in single rows; outer pores elongate, with similarly sized inner 
pores comma-shaped. Frontal sinus shallow. 

Etymology 

Ovatus L. egg- 
shaped, in reference to 
the form of the test, 
distinctive amongst the 
Schizasteridae. 

Material and locality 

Holotype ANU 
60653, a complete naked 
test from the vicinity of 
Cape Gazelle, New 
Britain, East New Britain 
Province, PNG. 



Description 

Test of small size, 
elongate ovoid, with 
length x width x height 
measuring 34 x 28 x 18 
mm; test length:width = 
1.21,width:height= 1.55. 
Test moderately 

depressed, with apical 
system located 55 percent 
of test length from 
anterior; test highest 
posterior to apical system, 
along keel of ambulacrum 
V. Oral surface is gently 




Figure 6. Schizaster (Paraster) ovatus sp. nov. Cape Gazelle, East New Britain. 
6a-d, ANU 60653, aboral, oral, lateral, posterior views. Bar scale = 2.5 mm. 



Proc. Linn. Soc. N.S.W., 125, 2004 



131 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 

convex. Apical system ethmolytic, depressed, with four genital pores, posterior pair being larger than anterior 
pair. Frontal ambulacrum long, shallow and narrow (12 percent of test length); pore pairs inclined at about 45° 
and arranged in a single row. Outer pores elongate, with similarly sized inner pores comma-shaped. Frontal 
sinus broad and shallow. Interambulacra II and III form sharp, high keels. Anterior petals diverging at angle of 
80°; flexed distally and shallow, bearing pore pairs which are elliptical, widely spaced and conjugate; 26 pairs 
are present. Posterior petals are moderately long (occupying 21 percent of test length), bearing 18 pore pairs. 

Peripetalous fasciole is distinct, passing transversely between posterior petals and thickening at petal 
ends; the fasciole describes a concave arc between the extremities of the posterior and anterior petals, with an 
outwards flexure, corresponding with a constriction, forward of the apical system. Fasciole reaches maximum 
thickness at the extremities of the anterior petals. Peripetalous fasciole passes forward from anterior petals at 
about 60° before curving strongly to close with frontal ambulacrum; constrictions occur on interambulacral 
keel and adjacent to the abrupt curvature. Lateroanal fasciole is narrower than peripetalous fasciole and of more 
constant width. Lateroanal fasciole extends abaxially posteriorly from peripetalous fasciole at constriction between 
posterior and anterior petals; at ambitus it runs far below periproct, close to adoral surface. 

Peristome oval and slightly sunken; situated anteriorly, anterior tip of labrum 15 percent of test length 
from anterior. Anteriorly labrum is strongly curved; bounded by thick rim that degenerates laterally. Labrum as 
long as broad; posterior extension triangular, about as long as broad. Labrum carries several small tubercles 
anteriorly. Plastron is pear-shaped and broad, maximum width being 3/4 length. Plastron tubercles are arranged 
in curving rows. 

Periproct at mid-level on sub-truncate end of test. Periproct longitudinally elliptical, with a prominent 
narrow slit extending a short distance axially and aborally towards interambulacrum V, nearly reaching apical 
surface (Fig. 6d). 

Remarks 

Schizaster (Paraster) ovatus sp. nov. can be distinguished from other Schizaster-like. heart urchins by its 
small, distinctively narrower and less inflated test, and long, shallow and narrow frontal ambulacrum. The test 
L: W and L:H ratios of 1 .2 1 and 1 .88 are larger than for most other echinoids of this group. The presence of four 
genital pores would suggest that the holotype is a mature specimen. McNamara and Philip (1980b) noted that in 
Schizaster (Ova) myorensis McNamara and Philip (1980b) the onset of maturity, occurring at a test length of 
about 25 mm, followed the sequential opening of the first, second, third and fouth genital pores. 

Morphological adaptations in Schizaster-like heart urchins are related to a need to produce a more efficient 
current flow over the aboral surface in sediment of low permeability (McNamara and Philip 1980a). The posterior 
migration of the apex meant more water would flow over over the frontal sinus to the peristome; the deepening 
of the frontal ambulacrum and the frontal sinus assisted in channelling water to the peristome; and the deep and 
long frontal ambulacrum further enabled more-funnel-building tube feet to be accommodated, presumably in 
response to finer-grained sediment (McNamara and Philip 1980a). The weakly vaulted test of S.(P.) ovatus 
with its shallow, open frontal ambulacrum and shallow frontal sinus suggests the species was a shallow-burrower 
in coarse (permeable) shell gravel. 

Suborder MICRASTERINA Fischer 1966 
Family BRISSIDAE Gray 1855 
Genus BRISSUS Gray 1825 

Synonymy 

Bryssus Martens 1869, p. 128 (nom. van.). 
Brissus (Allobrissus) Mortensen 1950, p. 162. 

Type species 

Spatangus brissus unicolour Leske 1778, p. 248 by subsequent designation of ICZN, Op. 290 1948. 

Brissus (Brissus) latecarinatus (Leske 1778) 

Synonymy 

Brissus carinatus Gray 1825, p. 431; A. Agassiz 1872-74, p. 96, 596. 

Brissus latecarinatus (Leske 1778): H.L. Clark 1921, p. 153; H.L. Clark 1925, p. 219; H.L. Clark 

132 Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



1946, p. 375; Mortensen 1951, p. 514; A.M. Clark and Rowe 1971, p. 165; Gibbs et al. 1976, 

p. 135. 
Brissus (Brissus) latecarinatus: Rowe and Gates 1995, p. 187. 
Spatangus Brissus latecarinatus Leske 1778, p. 249. 
Mortensen (1951: 514) lists additional synonymies. 

Material and locality 

Three naked tests, ANU 60643-5, from Nosnos village, Boang Island, Tanga Group, New Ireland 
Province, PNG; one naked test, B 20014, from Cape Gazelle, New Britain, East New Britain Province, PNG. 

Remarks 

Brissus (Brissus) latecarinatus (Leske 1778) is a widely distributed species throughout the Indo-Pacific 
(Mortensen 1951; A.M. Clark and Rowe 1971). It is present on Australian coasts, from Queensland to Port 
Jackson, and is also known from Lord Howe Island (H.L. Clark 1946). Miskelly's (2002) record of the species 
from the Solomon Islands is nearest to the present record in the Tanga Group. The largest specimen, ANU 
60644 from the Tanga Group, measures 70 x 60 x 39 mm, considerably smaller than the largest known specimen, 
from Hawaii, measuring 130 x 108 x 74 mm (HL. Clark 1946). The shape of the periproct of the Tanga Group 
and Cape Gazelle specimens, somewhat pointed above and below, differs from the rounded periproct evident in 
specimens figured by Mortensen (1951: Plate XXXHI, fig. 7) and Miskelly (2002). In this respect, the Bismarck 
Sea specimens closely resemble Brissus (Allobrissus) agassizii Doderlein 1885 (Mortensen 1951: Plate XXXJJI, 
fig. 7). Gibbs et al. (1976) noted the similarity of a Pelican Island, Great Barrier Reef, specimen of B. (B.) 
latecarinatus with B. (A.) agassizii. The posterior end of this particular specimen, like that of B. (A.) agassizii, 
is vertically truncated, with the posterior interambulacrum being only slightly carinate aborally (and not prolonged 
backwards to overhang the periproct and conceal it from dorsal view). 

Genus METALIA Gray 1855 

Synonymy 

Xanthobrissus Agassiz 1863, p. 28. 
Prometalia Pomel 1883, p. 34. 
Eobrissus Bell 1904, p. 236. 
Metaliopsis Fourtau 1913, p. 68. 

Type species 

Spatangus sternalis Lamarck 1816, p. 326, by original designation. 

Metalia spatagus (Linnaeus 1758) 

Synonymy 

Echinus spatagus Linnaeus 1758, p. 665. 

Metalia spatagus (Linnaeus 1758): H.L. Clark 1925, p. 216; H.L. Clark 1932, p. 219; HL. Clark 

1946, p. 372; Mortensen 1951, p. 540; A.M. Clark and Rowe 1971, p. 166; Gibbs et al. 1976, 

p. 136; Rowe and Gates 1995, p. 190. 
Mortensen (1951: 540) lists additional synonymies. 

Material and locality 

Two naked tests, ANU 60646-7, from Nosnos village, Boang Island, Tanga Group, New Ireland 
Province, PNG. 

Remarks 

Metalia spatagus (Linnaeus 1758) is widely distributed through the Indo-Pacific (Mortensen 1951; A.M. 
Clark and Rowe 1971). H.L. Clark (1932) provided the first record of this species from Australasian waters 
(Low Isles, Great Barrier Reef), recording the largest known specimen, measuring 1 10 x 93 x 52 mm. By 
comparison, the largest Tanga specimen measures 54 x 40 x 29 mm. Miskelly (2002) records the species from 
the Solomon Islands. 



Proc. Linn. Soc. N.S.W., 125, 2004 133 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 



ACKNOWLEDGMENTS 

The author is grateful to Alistair Norrie and 
Richard Joycey, East New Britain Historical and Cultural 
Centre, for the loan of specimens from the Kokopo 
Museum, Kokopo, PNG. Prof. Ken Campbell kindly 
provided his thoughts on the erection of new species 
Schizaster (Paraster) ovatus sp. nov. and Heliocidaris 
robertsi sp. nov. and Loisette Marsh, Western Australian 
Museum, kindly provided an opinion on the identification 
of the juvenile Protoreaster nodosus from New Britain. 
Dr. Richard Barwick and Dr. Make LeGleuher, both of 
the Department of Geology, Australian National 
University, kindly photographed all specimens, and 
provided translation from French of sections from Koehler 
(1910) and De Ridder (1986), respectively. The comments 
of Geoff Francis and an anonymous reviewer improved 
the manuscript. 



REFERENCES 

Agassiz, A. (1863). List of the echinoderms sent to 
different institutions in exchange for other 
specimens, with annotations. Bulletin of the 
Museum Comparative Zoology, Harvard 1(2) 
17-28. 

Agassiz, A. (1879). Preliminary report on the Echini 
of the exploring expedition of H.M.S. 
Challenger. Proceedings of the American 
Academy of Arts and Sciences 14 190-212. 

Agassiz, A. (1881). Report on the scientific results of 
the voyage of H.M.S. Challenger during the 
years 1873-76: Zoology - vol. 3, pt. 9, Report 
on the Echinoidea. 321pp. (H.M. Stationery 
Office: London). 

Bell, F.J. (1899). Report on the Echinoderms (other 
than Holothurians) collected by Dr. Willey. 
In 'Zoological results based on material from 
New Britain, New Guinea, Loyalty Islands 
and elsewhere, collected during the years 
1895 1896 and 1897' (Ed. A. Willey) Part II 
133-140. (Cambridge University Press: 
Cambridge). 

Bouillon, J. and Jangoux, M. (1984). Note on the 
relationship between the parasitic mollusk 
Thyca crystallina (Gastropoda, 
Prosobranchia) and the starfish Linckia 
laevigata (Echinodermata) on Laing Island 
reef (Papua New Guinea). Annales de la 
Societe Roy ale Zoologique de Belgique 114, 
249-256. 

Britayev, T.A., Doignon, G. and Eeckhaut, I. (1999). 
Symbiotic polychaetes from Papua New 
Guinea associated with descriptions of three 
new species. Cahiers de Biologie Marine 40, 
359-374. 

Chapman, F. and Cudmore, F.A. (1934). The 

Cainozoic Cidaridae of Australia. Memoirs of 
the National Museum Melbourne 8 126-149. 



Clark, A.H. (1954). Records of Indo-Pacific 

echinoderms. Pacific Science 8, 243-263. 

Clark, A.M. (1968). 'Starfishes and their relations'. 
(British Museum (Natural History): London). 
120pp. 

Clark, A.M. and Rowe, F.W.E. (1971). Monograph 
of shallow-water Indo-West Pacific 
echinoderms. British Museum (Natural 
History) Publication 690, 238pp. 

Clark, HL. (1908). Some Japanese and East Indian 
echinoderms. Bulletin of the Museum of 
Comparative Zoology, Harvard 51 (11), 279- 
311. 

Clark, HL. (1925). A catalogue of the recent Sea- 
Urchins (Echinoidea) in the collection of the 
British Museum (Natural History). (British 
Museum (Natural History): London). 250pp. 

Clark, HL. (1932). Echinodermata (other than 

Asteroidea). Scientific Reports of the Great 
Barrier Reef Expedition 4 197-239. 

Clark, HL. (1946). The Echinoderm Fauna of 

Australia: Its Composition and Its Origin. 
Carnegie Institution of Washington, 
Publication 566, 567pp. 

Corbett, G., First, D.M. and Hayward, S.B. (1991). 
The Maragorik Prospect, east New Britain, 
Papua New Guinea. In 'Proceedings of the 
PNG Geology, Exploration and Mining 
Conference 1991, Rabaul' (Ed. R. Rogerson) 
pp. 112-116. (The Australasian Institute of 
Mining and Metallurgy: Melbourne). 

De Ridder, C. (1986). Les echinides. In 'Guide des etoiles 
de mer, oursins et autres echinodermes du lagon 
de Nouvelle-Caledonie' (Eds A. Guille, P. 
Laboute and J.-L. Menou) pp. 22-53. (Institut 
Francais de Recherche Scientifique pour le 
Developpement en Cooperation. Collection Faune 
Tropicale 25). 

Durham, J.W. (1966). Phylogeny and evolution. In 
'Treatise on Invertebrate Paleontology, Part 
U, Echinodermata 3' (Ed. R.C. Moore) pp. 
U266-U269. (Geological Society of America 
and University of Kansas Press: Lawrence). 

Eeckhaut, I., Deheyn, D. and Jangoux, M. (1996). 
Study on the symbiotic fauna of crinoids 
collected in Hansa Bay (Bismarck Sea, Papua 
New Guinea). Ninth International 
Echinoderm Conference, San Francisco, 
August 1996. pp. 516. (A.A. Balkema, 
Rotterdam). 

Endean, R. (1957). The biogeography of Queensland's 
shallow-water echinoderm fauna (excluding 
Crinoidea), with a rearrangement of the faunistic 
provinces of tropical Australia. Australian Journal 
of Marine and Freshwater Research 8, 233-273. 

Fell, H.B. (1966). Cidaroids. In 'Treatise on 
Invertebrate Paleontology, Part U, 
Echinodermata 3' (Ed. R.C. Moore) pp. 
U3 1 2-U340. (Geological Society of America 
and University of Kansas Press: Lawrence). 



134 



Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



Fell, H.B. and Pawson, D.L. (1966). Echinacea. In 
'Treatise on Invertebrate Paleontology, Part 
U, Echinodermata 3' (Ed. R.C. Moore) pp. 
U367-U440. (Geological Society of America 
and University of Kansas Press: Lawrence). 

Fischer, A.G. (1966). Spatangoids. In 'Treatise on 

Invertebrate Paleontology, Part U, Echinodermata 
3' (Ed. R.C. Moore) pp. U543-U628. (Geological 
Society of America and University of Kansas 
Press: Lawrence). 

Fisher, W.K. (1919). Starfishes of the Philippine 
Seas and adjacent waters. United States 
National Museum Bulletin 100(3), 711pp. 

Francis, G. (1985). Stratigraphy of Manus Island, 
western New Ireland Basin, Papua New 
Guinea. Geological Survey of Papua New 
Guinea Report 85/10, 40 pp. 

Gibbs, P.E., Clark, A.M. and Clark, CM. (1976). 

Echinoderms from the northern region of the 
Great Barrier Reef, Australia. Bulletin of the 
British Museum (Natural History) (Zoology) 
30(4) 103-144. 

Koehler, R. (1910). An account of the shallow-water 
asteroidea. Echinoderma of the Indian 
Museum. (Trustees of the Indian Museum: 
Calcutta). 192pp. 

Lindley, I.D. (1988). Early Cainozoic stratigraphy 
and structure of the Gazelle Peninsula, east 
New Britain: An example of extensional 
tectonics in the New Britain arc-trench 
complex. Australian Journal of Earth 
Sciences 35, 231-244. 

Lindley, I.D. (2003a). Echinoids of the Kairuku 

Formation (Lower Pliocene), Yule Island, Papua 
New Guinea: Clypeasteroida. Proceedings of the 
Linnean Society of New South Wales Y2A 125- 
136. 

Lindley, I.D. (2003b). Echinoids of the Kairuku 

Formation (Lower Pliocene), Yule Island, Papua 
New Guinea: Regularia. Proceedings of the 
Linnean Society of New South Wales 124 137- 
151. 

Loriol, P. de. (1891). Notes pour servir a l'etude des 
Echinoderms. III. Mem. Soc. Phys. Hist. not. 
Geneve vol. suppl. 8 1-31. 

Martin, K. (1918). Unsere palaeozooligische Kentiss von 
Java. Beilage-Band zu Sammlungen des 
Geologischen Reischsmuseums in Leiden. 118pp. 

McNamara, K.J. and Kendrick, G.W. (1994). Cenozoic 

molluscs and echinoids of Barrow Island, Western 
Australia. Records of the Western Australian 
Museum, Supplement No. 51, 50pp. 

McNamara, K.J. and Philip, G.M. (1980a). Australian 

Tertiary schizasterid echinoids. Alcheringa 4, 47- 
65. 

McNamara, K.J. and Philip, G.M. (1980b). Living 

Australian schizasterid echinoids. Proceedings of 
the Linnean Society of New South Wales 104 127- 
146. 

Messing, C.G. (1994). Comatulid crinoids 



(Echinodermata) of Madang, Papua New Guinea 

and environs: diversity and ecology. In 

'Echinoderms through time' (Eds D.B. Guille, A.. 

Feral, J.-P. and M. Roux) pp. 237-243. (A.A. 

Balkema: Rotterdam). 
Miskelly, A. (2002). 'Sea urchins of Australia and the 

Indo-Pacific'. (Capricornica Publications: 

Sydney). 179 pp. 
Mortensen, T. (1928a). A Monograph of the Echinoidea I, 

Cidaroidea. (C.A. Reitzel, Copenhagen). 551 pp. 
Mortensen, T. (1928b). Papers from Dr. Th. Mortensen's 

Pacific expedition 1914-16. New Cidaridae. 

Videnskabelige Meddelelser fra Dansk 

Naturhistorisk Forening 1 Ktfkenhavn 85, 65-74. 
Mortensen, T. (1943a). A Monograph of the Echinoidea 

111.2 Camarodonta I: Orthopsidae, 
Glyphocyphidae, Temnopleuridae and 
Toxopneustidae. (C.A. Reitzel, Copenhagen). 553 
pp. 

Mortensen, T. (1943b). A Monograph of the Echinoidea 

111.3 Camarodonta II: Echinidae, 
Strongylocentrotidae, Parasalenidae, 
Echinometridae. (C.A. Reitzel, Copenhagen). 446 
pp. 

Mortensen, T. (1948a). A Monograph of the Echinoidea 
IV. 1 Holectypoida, Cassiduloida. (C.A Reitzel, 
Copenhagen). 371 pp. 

Mortensen, T. (1948b). A Monograph of the Echinoidea 
IV.2 Clypeasteroida: Clypeasteridae, Arachnoidae, 
Fibulariidae, Laganidae and Scutellidae. (C.A. 
Reitzel, Copenhagen). 471 pp. 

Mortensen, T. (1951). A Monograph of the Echinoidea 
V.2 Spatangoida II: Amphisternata II, 
Spatangidae, Loveniidae, Pericosmidae, 
Schizasteridae, Brissidae. (C.A. Reitzel, 
Copenhagen). 593 pp. 

Miiller, J. and Troschel, F.H. (1842). 'System der 
Asteriden'. (Papier, Druck and Verlag: 
Braunschweig). 134 pp. 

Negretti, B., Philippe, M., Soudet, H.J., Thomassin, BA. 
and Oggiano, G. (1990). Echinometra miocenica 
Loriol, echinide Miocene, synonyme 
d' Echinometra mathaei (Blainville), actuel: 
Biogeographie et paleoecologie. Geobios 23, 445- 
456. 

Philip, G.M. (1965). The Tertiary echinoids of south- 
eastern Australia III Stirodonta, Aulodonta, and 
Camarodonta (1). Proceedings of the Royal 
Society of Victoria 78(2): 181-196. 

Read, J.R.L. (1965). Preliminary geological investigation 
of the lower Warangoi hydro-electric scheme. 
New Britain August-November 1964. Bureau of 
Mineral Resources, Geology and Geophysics, 
Record 1982/15. 

Rowe, F.W.E. and Gates, J. (1995). Echinodermata. In 

'Zoological Catalogue of Australia', Vol. 33. 510 
pp. (CSIRO Australia: Melbourne). 

Sladen, W.P. (1889). Report on the scientific results of the 
voyage of H. M.S. Challenger during the years 
1873-76: Zoology - vol. 30, Report on the 



Proc. Linn. Soc. N.S.W., 125, 2004 



135 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 



Asteroidea. 893pp. (H.M. Stationery Office: 
London). 

Spencer, W.K. and Wright, C.W. (1966). 

Asterozoans. In 'Treatise on Invertebrate 
Paleontology, Part U, Echinodermata 3' (Ed. 
R.C. Moore) pp. U4-107. (Geological Society 
of America and University of Kansas Press: 
Lawrence). 

Sluiter, C.P. (1895). Die Asteriden Sammlung des 
Museums zu Amsterdam. Die Echiniden 
Sammlung des Museums zu Amsterdam. 
Bijdragen tot de Dierkunde 17, 49-64, 65-74. 

Struder, T. (1876). Uber Echinodermen aus dem 

antarktischen Meere und zwei neue Seeigel von 
den Papua-Inseln gesammelt auf der Reise S.M.S. 
'Gazelle' um die Erde. Monatsberichte der 
Koniglich Preussischen Akademie der 
Wissenschaften zu Berlin 1876, 452-465. 

Struder, T. (1880). Ubersicht iiber die wahrend der Reise 
S.M.S. Corvette 'Gazelle' um die Erde 1874-76 
gesammelten Echinoiden. Monatsberichte der 
Koniglich Preussischen Akademie der 
Wissenschaften zu Berlin 1880, 861-885. 

Wagner, CD. and Durham, J.W. (1966). Holectypoids. In 
'Treatise on Invertebrate Paleontology, Part U, 
Echinodermata 3' (Ed. R.C. Moore) pp. U440- 
450. (Geological Society of America and 
University of Kansas Press: Lawrence). 

Wallace, D.A., Johnson, R.W., Chappell, B.W., Arculus, 
R.J., Perfit, M.R. and Crick, I.H. (1983). 
Cainozoic volcanism of the Tabar, Lihir, Tanga 
and Feni Islands, Papua New Guinea: Geology, 
whole-rock analyses, and rock-forming mineral 
compositions. Bureau of Mineral Resources, 
Geology and Geophysics, Report 243. 

Willey, A. (1902). Zoological results based on material 
from New Britain, New Guinea, Loyalty Islands 
and elsewhere, collected during the years 1 895 
1896 and 1897. Pt. 2. (Cambridge University 
Press: Cambridge). 253pp. 

Woods, H. (1958). 'Palaeontology Invertebrate'. 

(Cambridge University Press; Cambridge). 477pp. 



136 Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 

Table 1. Reported starfishes from the Bismarck Archipelago, Papua New Guinea. 

ASTERENAE 

Tarsaster stoichodes Sladen 1889: Fisher 1919, p. 491: north of the Admiralty Group (150 fathoms). 

ASTEROID AE 

Asterina cephus (Muller and Troschel 1842): A.H. Clark 1954, p. 258: Seleo Island, Aitape district. 
Patiriella exigua (Lamarck 1816): A.H. Clark 1954, p. 258: Admiralty Group; Seleo Island, Aitape district. 

ASTEROPSEIDAE 

Asteropsis carinifera (Lamarck 1816): A.H. Clark 1954, p. 258: Seleo Island, Aitape district. 

ASTROPECTINHDAE 

Astropecten monacanthus Sladen 1883: Bell 1899, p. 136: New Britain. 

Astropecten polyacanthus Muller and Troschel 1842: Fisher 1919, p. 64: Admiralty Group. 

ECHTNASTERIDAE 

Echinaster luzonicus (Gray 1840): Rowe and Gates 1995, p. 59. (= Echinaster eridanella Muller and 
Troschel 1842, p. 24; Bell 1899, p. 138): New Ireland; New Britain. 

LUIDIIDAE 

*Luida aspera Sladen 1889: Fisher 1919, p. 171: north of Admiralty Group (150 fathoms). 

OPHIDIASTEPJDAE 

Linckia laevigata (Linnaeus 1758): Bouillon and Jangoux 1984, p. 249: Laing Island reef, Hansa Bay. 
Nardoa novaecaledoniae (Perrier 1875): Rowe and Gates 1995, p. 88. (= Nardoa mollis de Loriol, 1891, 

H.L. Clark 1946, p. 115; A.H. Clark 1954, p. 255): New Britain; Seleo Island, Aitape district. 
Nardoa tuberculata Gray 1840: Rowe and Gates 1995, p. 88. (= Nardoa finschi de Loriol 1891; Nardoa 

pauciforis von Martens 1866, H.L. Clark 1946, p. 115): New Britain. 
Ophidiaster granifer Liitken 1871: A.H. Clark 1954, p. 256: Seleo Island, Aitape district. 

OREASTERIDAE 

+Anthenea sidneyensis Doderlein 1915: Rowe and Gates 1995, p. 98: Manus Island (Admiralty Group). 
Culcita novaeguineae Muller and Troschel 1842: A.H. Clark 1954, p. 254: Seleo Island, Aitape district. 
Pentaster obtusatus (Bory de St. Vincent 1827). [= Pentaceropsis obtusata (Bory de St. Vincent 1827) Bell 

1899, p. 136]: Blanche Bay, New Britain. 
Protoreaster lincki (de Blainville 1830): Oreaster lincki (= Pentaceros lincki, Bell 1899, p. 136): Blanche 

Bay, New Britain. 
Protoreaster nodosus (Linnaeus 1758): H.L. Clark 1946, p. 106; A.H. Clark 1954, p. 254. (= Pentaceros 

nodosus, Bell 1899: p. 136; Oreaster nodosus H.L. Clark 1908): Blanche Bay, New Britain; Seleo 

Island, Aitape district. 

PTERASTERIDAE 

Hymenaster pullatus Sladen 1889: Fisher 1919, p. 467: southwest of the Admiralty Group (1,070 fathoms). 

NOTES 

+ the writer follows Spencer and Wright (1966) and Rowe and Gates (1995) in placing Anthenea in Family 
Oreasteridae. H.L. Clark (1946) and A.M. Clark and Rowe (1971) placed the taxon in Family Goniasteridae. 

* Denotes type locality in Bismarck Archipelago. 



Proc. Linn. Soc. N.S.W., 125, 2004 137 



FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 

Table 2. Reported shallow and deep-water sea-urchins from the Bismarck Archipelago, Papua New 
Guinea. 

ARACHNOIDIDAE 

A rachnoides placenta (Linnaeus 1758): Bell 1899, p. 136; H.L. Clark 1925, p. 154: New Britain. 

ARBACIIDAE 

*Pygmaeocidaris prionigera (A. Agassiz 1879): A. Agassiz 1881, pi. XXXTV, figs 14 and 15; H.L. Clark 

1925, p. 73 (= Podocidaris prionigera A. Agassiz 1879, p. 199): between New Guinea and Admiralty 

Group (1,070 fathoms). 
*Coelopleurus elegans (Bell 1899): H.L. Clark 1925, p. 73. (= Salmacis elegans Bell 1899, p. 135): New 

Britain. 

CIDARIDAE 

Eucidaris metularia (Lamarck 1816): H.L. Clark 1925, p. 20. (= Cidaris metularia de Blainville, 1830, Bell 

1899, p. 134): New Britain. 
Prionocidaris baculosa var. annulifera (Lamarck): Mortensen 1928a, p. 437, 446. (= Schleinitzia crenularis 

Struder 1876, p. 463; 1880, p. 865): west New Guinea. 
Stylocidaris reini (Doderlein): H.L. Clark 1925, p. 24; Mortensen 1928a, p. 342, 347, 474 (= Phyllacanthus 

annulifera Bell 1899, p. 134): New Britain; Milne Bay. 

DIADEMATIDAE 

Echinothrix calamaris (Pallas 1774): A.H. Clark 1954, p. 250: Bougainville Island. 

*Micropyga nigra H.L. Clark 1925: A. Agassiz 1879, p. 200; H.L. Clark 1925, p. 47. (= Astropyga elastica 

Struder, Bell 1899, p. 135): New Britain. 
Micropyga tuberculata A. Agassiz 1879, p. 200: A. Agassiz 1881, pi. VII; H.L. Clark 1925, p. 48: Blanche 

Bay, New Britain. 

ECHINOMETRIDAE 

Echinometra mathaei (de Blainville 1825): A.H. Clark 1954, p. 251: Bougainville Island; Seleo Island, 
Aitape district; Normanby Island. (= Echinometra lucunter Bell 1899, p. 136). 

ECHINOTHURIIDAE 

Araeosoma gracile (A. Agassiz 1881): A. Agassiz 1881, p. 89; H.L. Clark 1925, p. 61: Admiralty Group 
(150 fathoms). 

LAGANIDAE 

Laganum decagonale (de Blainville 1827): A. Agassiz 1881; H.L. Clark 1925, p. 156; Mortensen 1948b, p. 

332, 336; Lindley 2003a, p. 133: near Admiralty Group (150 fathoms). 
Laganum depressum var. tonganense (Quoy and Gainard): Mortensen 1948b, p. 324: Admiralty Group. 
Laganum laganum (Leske): Mortensen (1948b), p. 312: Bismarck Archipelago. 

SPATANGIDAE 

Maretia ovata (Leske 1778): A. Agassiz 1881; H.L. Clark 1925, p. 226: Admiralty Group. 

TEMNOPLEURIDAE 

Prionechinus agassizii Wood-Mason and Alcock 1891: H.L. Clark 1925, p. 78. (= Echinus elegans, A. 

Agassiz 1881): near Admiralty Group. 
* Prionechinus sagittiger A. Agassiz 1879, p. 202: A. Agassiz 1881, pi. IVa, figs 11-14; H.L. Clark 1925, p. 

79: between New Guinea and Admiralty Group (1,070 fathoms). 
Temnopleurus sp., Bell 1899, p. 135: New Britain. 
Temnopleurus reevesii (Gray 1855): A. Agassiz 1881; H.L. Clark 1925, p. 81: near Admiralty Group (150 

fathoms). 



138 Proc. Linn. Soc. N.S.W., 125, 2004 



I.D. LINDLEY 



Temnotrema scillae (Mazetti 1894): Mortensen 1904, p. 86; H.L. Clark 1925, p. 91 (= Pleurechinus 
reticulatus in H.L. Clark 1925, p. 91): New Britain. 

TOXOPNEUSTIDAE 

Tripneustes gratilla (Linnaeus 1758): A.H. Clark 1954, p. 250: Bougainville Island; Seleo Island, Aitape 
district. 

INVALID RECORDS 

Astriclypeus manni Verrill, Sluiter 1895, p. 73, New Ireland; Mortensen 1948b, p. 416, 418. 
Colobocentrotus mertensi Brandt 1835, Sluiter 1895, p. 69, New Ireland; Mortensen 1943b, p. 433. 
Mellita longifissa Michelin 1858, Sluiter 1895, p. 73, New Ireland; Mortensen 1948b, p. 427, 428. 
Taxonomic reason: Erroneous labelling (Mortensen 1948b, p. 418; Mortensen 1943b, p. 433; Mortensen 
1948b, p. 428, respectively). 



NOTES 

* Denotes type localities in Bismarck Archipelago 



Proc. Linn. Soc. N.S.W., 125, 2004 139 



140 



Conodont Faunas from the Mid to Late Ordovician Boundary 

Interval of the Wahringa Limestone Member (Fairbridge 

Volcanics), Central New South Wales 

Y.Y. Zhen 1 *, I.G. Percival 2 *and B.D. Webby 3 

'Division of Earth and Environmental Sciences, The Australian Museum, 6 College Street, Sydney, N.S.W. 

2010 (yongyi@austmus.gov.au); 2 Geological Survey of New South Wales, Department of Mineral 

Resources, P.O. Box 76, Lidcombe, N.S.W. 2141 (ian.percival@minerals.nsw.gov.au); 3 Centre for 

Ecostratigraphy and Paleobiology, Department of Earth and Planetary Sciences, Macquarie University, 

N.S.W. 2109, Australia (bwebby@laurel.ocs.mq.edu.au); *Honorary Research Associate, Centre for 

Ecostratigraphy and Paleobiology, Macquarie University, N.S.W. 



Zhen, Y.Y., Percival, I.G. and Webby, B.D. (2004). Conodont faunas from the Mid to Late Ordovician 
boundary interval of the Wahringa Limestone Member (Fairbridge Volcanics), central New South 
Wales. Proceedings of the Linnean Society of New South Wales 125, 141-164. 

Twenty-nine conodont species are documented from the Wahringa Limestone Member and other isolated 
limestone pods of the Fairbridge Volcanics, in the Bakers Swamp area between Wellington and Orange, 
central New South Wales. Three conodont assemblages are recognised within the Wahringa Limestone 
Member. The oldest is characterised by the occurrence of Pygodus protoanserinus and Pygodus serra, 
indicative of a late Darriwilian age (Da3 to early Da4). The overlying assemblage B, bearing Belodina 
monitorensis, probably ranges across the Mid to Late Ordovician boundary. Assemblage C with abundant 
Belodina compressa in the upper part of the Wahringa Limestone Member is of late Gisbornian (Gi2) age. 
The conodont faunas are significant in being the first described from the Lachlan Orogen in New South 
Wales spanning the Mid to Late Ordovician interval, although resolution of the actual boundary level is 
limited in the section measured. 

Manuscript received 17 September 2003, accepted for publication 17 December 2003. 

KEYWORDS: Conodonts, Fairbridge Volcanics, Late Ordovician (Gisbomian), Mid Ordovician 
(Darriwilian), Wahringa Limestone Member. 



report defining this unit, and their age connotations 

INTRODUCTION discussed (Percival et al. 1999). Subsequent detailed 

sampling has yielded many more elements and species, 

The Molong Volcanic Belt (MVB) is a enabling broad confirmation of the original age 

meridionally-aligned tectonic feature of Ordovician determination and providing increased precision for 

age within the east Lachlan Orogen in central New the upper age limit of the ii mes tone. The faunas are 

South Wales (Glen et al. 1998). Ordovician strata in systematically described here for the first time. They 

the northern sector of the MVB near Bakers Swamp we significant in spanning the Mid to Late Ordovician 

between Orange and Wellington (Fig. 1) are boundary, an interval which is otherwise poorly 

represented by the Early Ordovician Mitchell represented in shallow water settings of eastern parts 

Formation and Hensleigh Siltstone - the latter yielding f trie Lachlan Orogen. 
conodonts of Bendigonian age (upper Prioniodus 
elegans Zone) from allochthonous limestones (Zhen 

et al. in press) - and the Fairbridge Volcanics of Mid STRATIGRAPHY 
to early Late Ordovician age. Two autochthonous 

limestone units occur in the Fairbridge Volcanics. The Much of the Fairbridge Volcanics consists of 

lower one, known as the Wahringa Limestone an desitic lava flows, with subsidiary boulder 

Member, is the subject of this paper. Representative conglomerates (Morgan, Scott & Percival, in Meakin 

conodonts were illustrated earlier in a preliminary & Morgan 1999). Allochthonous limestones are 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 




CATOMBAL 
GROUP 



C1474, C1483 

C1472 

C1695-C1700 

C1471.C1693, C1694 

C1429 
WAHRINGA C1458 

LIMESTONE 
MEMBER C1456 

C1450 '- 
C1486, C1487, C1488 
C1463 



'CZX 



HENSLEIGH 
SILTSTONE 



MITCHELL 
FORMATION 



LOCATION MAPS 



-/-*-? fl 






\ 



AUSTRALIA 



X) 

^7 

NEW SOUTH WALES / 

Wellington/ 
Study Area-* J 



cd 2 c 

TJ J3 •- 

S>- > 
!s ° 

» S "s 

«oO 



REFERENCE 
I Quaternary alluvium 
_J Undifferentiated Devonian 
\ A * m \ Undifferentiated Silurian 

| A | Fairbridge Volcanics 

allochthono 
limestones 



Dip, and strike 
_». Syncline, with plunge 
_ Anticline 



□ allochthonous 
lin 



I Wahringa Limestone 
I Member 



Orange * 



7 Sydney 



M 



~J Hensleigh Siltstone 
[■Jjl] limestones 
| -j J Mitchell Formation 



C1458. Fossil locality 

*■ Thrust fault 
===== Road 

• Type section 



1 



2 km 

i 



Figure 1. Locality maps. A. location of Wahringa area, between Wellington and Orange, central New 
South Wales; B. simplified geological map of the Wahringa area; C. generalised stratigraphic column for 
this area, showing spot sampled horizons within the Wahringa Limestone Member and the allochthonous 
limestone blocks within the Fairbridge Volcanics. For further details of the sampled section, see Figure 2. 



142 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



uncommon in the lower Fairbridge Volcanics below 
the Wahringa Limestone Member and conodont yields 
are disappointingly low. Numerous allochthonous 
limestone pods emplaced within the Fairbridge 
Volcanics, stratigraphically overlying the Wahringa 
Limestone Member, were also processed for 
conodonts. With the exception of six samples that 
contained Belodina compressa (indicating an early 
Late Ordovician or Gisbornian age), these were either 
barren or yielded only sparse, non-diagnostic elements. 

Wahringa Limestone Member of Fairbridge 
Volcanics (Fig. 2) 

The name derives from the "Wahringa" 
property, located approximately 28 km south of 
Wellington on the Mitchell Highway. Here the 
Wahringa Limestone Member is exposed along strike 
for approximately 400 m and attains a thickness of 88 
m in its type section (situated just north of a bend in 
Bakers Swamp Creek). Invertebrate macrofossils 
described or illustrated from the Wahringa Limestone 
Member comprise brachiopods, gastropods, nautiloids, 
crinoids, demosponges, stromatoporoids, and a species 
of tabulate coral (Percival et al. 2001). The unit is 
subdivisible into three parts: lower beds rich in 
oncolites, ooids and volcaniclastic detritus, a middle 



C1687 
C1683-J: 

to 
C1678 
C1677_ 
C1676£ 
C1675# 



C16747 
C1673' 



C1672- 




C1668- 
C1667" 

C1664- 



C1652- 
(C1450) 




3 

■Scv. 

fe § 
8 I 

ca 5: 

O <B 

i* 

So 

■o .y 
II 

• • 



§ * 



•j. assemblage C 

<D 
O 



h 

IB 8 

■S s- 
u & 

■a s 
<8 4 



assemblage B 



assemblage A 



r10m 



Figure 2. Stratigraphic section measured through 
the Wahringa Limestone Member showing sampled 
horizons and ranges of selected, age-significant 
conodont taxa. 



part of muddy, thinly bedded limestones rich in 
brachiopods, and an upper section that is more massive. 
The variation in lithologies reflects an increase in water 
depth from shallow subtidal at the base, to below 
normal wave base in the middle and upper beds. 
However, this depth increase does not have a 
controlling influence on the three distinct conodont 
assemblages recognised, which represent age- 
significant rather than biofacies-distinct assemblages. 

Lithologies in the lower part of the member, 
which consist mainly of red ooidal grainstones, 
calcarenites and oncolitic grainstone-packs tones, are 
particularly characteristic of very shallow deposition. 
Fauna present in these beds (not observed at other 
levels of the measured section) include the large 
gastropod Maclurites cf. M. florentinensis, the 
siphonotretoidean brachiopod Multispinula, and a 
Calathium-like receptaculitid. Demosponges, 
including A rchaeoscyphia! sp. B, Malongullospongial 
sp. and Hindia cf. H. sphaeroidalis, are more widely 
distributed but are especially common at this level. 
Fossil grains are subangular to rounded, frequently 
algal-coated and include dasycladacean and 
solenoporid algae. Large oncolites with well-preserved 
cyanobacterial Girvanella filaments and ooids are also 
abundant. 

The middle part of the Wahringa Limestone 
Member is characterised by fine to coarse grained 
skeletal grainstone (interbedded with silty layers) that 
is dominated by remains of echinoderms, brachiopods, 
dasycladacean algae, molluscs, trilobites, and 
ostracods. Brachiopods, including Sowerbyitesl, 
Leptellina and rare Sowerbyella are concentrated in 
thin-bedded packstones in the middle part of the unit. 
Stromatoporoids (Labechia, Labechiella), mostly 
preserved in growth position, occur slightly above this 
level and range into the uppermost limestone beds of 
the unit. 

The most common lithologies in the upper 
part of the Wahringa Limestone Member are fine to 
coarse grained skeletal, oncolitic grainstone and lesser 
packstone to wackestone lacking internal lamination. 
Grains include ostracods, dasycladacean algae, rare 
ooids, and oncolites with associated Girvanella. 



CONODONT BIOSTRATIGRAPHY 

Twenty-nine conodont species based on 897 
individual specimens were recovered from 44 samples 
(Fig. 3), collected from the Wahringa Limestone 
Member and various limestone pods within the 
enclosing Fairbridge Volcanics. The faunas range in 
age from late Darriwilian (Da3, lower Eoplacognathus 
suecicus Zone) to late Gisbornian (Gi2, Belodina 



Proc. Linn. Soc. N.S.W., 125, 2004 



143 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 



~ p -• 

^o 2 

•• cl" oj 

i.™ b 

& < K 

ST — 2. 

© 65 g 

1 3 # g. 
C o" 0* 

5 " B 

"» o 85 

o" 3 2. 

3- 65 O 

M> ° 2 

i 5 ^ § 

I § a 

M g o 

3 © 2 

« 3 2. 

1 « i 5 

i 3" r 

2 2 

3 3' 1/5 
3 3k 

SW 3 
^ B | 

o 3 2. 

EL « cr 

2 ° 2 
033 

« 5? ST 

l "* 5? & 

-r 3 » 

- if 



3 



3 
cro 



r 
3 1 



re 

"1 

o| 

2. era 

3 " 

3 

3 



3 
_ n 

«-• a) 

O 
3 
a 



2 » 

3 v> 
re £■ 



1^3 

-I 12. 65 

' 2> 3 

w a 

o SL 

n S3 

o 
3 « 

1 ST 

"* Si 

re sr 

3 O 

2 3 

o 3; 



c 
re 

ST 



3. ST 
re re 



H 
O 


1 
1 

i 

■8 

03 


I 

| 
> 


t 

s 

3 


1 
| 

3. 

5 


1 

1 

3 
i 


i 

1 
1 

•5' 

5 


1 

I 

1 

1 
1 


1 

1 

s 

1 

2 


I 

1 

! 

a 
| 


I 

l 

s 
1 

a. 


5 

i 
1 

i 

5 




1 

3 
1 

s 

1 




1 
s 


a 1 

I 

a 


I 

1 

a 
f 

•>3 


9 
1 

■3 


1 

1 

1 


I 

1 


1 


i 
1 
s 
■S 
m 


to 

I 
I 

•S 

> 


CD 
1 

3 
i 

I 

1 


| 

1 
1 


t 

i 

a 

1 

5 

1 

i 


1 
1 


8 

t 


1^ 
1 

a 


1 

■S 


O / 

3 / 

/ 

1 

3 / 

1 / 
B> / 

/ 0) 

/ >» 

/ 3 

/ "° 
/ 

/ w 


B 1 

i 

0. 
s 


8 




to 














tJ 






■^ 






- 




















- 




- 






£9H3 


D* 5" 


Z 


























- 




























- 






98W3 


S 


























Jfc 






















- 












i8MD 


I 
















































- 












88H3 


9£ 








- 










- 




- 


h- 





N» 




- 


- 




- 




- 














*. 


M 


OSt-13 


1 

3* 
1 

3 
(Q 

a 

I - 

3 


(A 
O 

3 
O 

3 


3 

O" 



J? 

■0 

ID 
CO 

CD 
O 

O 

3 


£1 
























« 


Xk 


to 


















*k 














9SHD 


6ZI 












- 




M 




w 




a 


a 










■u 










i- 


to 








- 




8SfI3 


£ 


























to 


- 
































WHO 


6S 






- 


\« 






- 










no 


a 








- 




- 




to 










- 






- 


ZS9I3 


fr 


























w 




















- 














W9I3 


£ 
























- 


to 


































i99I3 


Z 
























































u 




89913 


I 






















































- 






ZCSID 


fr 
























- 


- 


- 


















- 














£L910 


£T 















































b) 














W9I3 


OS 


N> 












-J 










to 


-j 


- 








w 


M 


*. 






W 


M 












SL9X3 


zz 
























w 


o\ 














- 








- 








- 




9L910 


£ 


























** 






















to 












LL913 


S 


























w 






















M 












8i9I3 


£ 










■ 
















- 




















• >3 


- 












6/.9T3 


9 
















- 










w 






















- 










- 


08913 


I 
















































- 












T89I3 


£1 




















w 




t*> 


ON 














- 








Ul 












Z89I3 


£1 




















- 




- 


Ul 






















Ut 








- 




£8913 


zz 
























w 


ON 




















to 











- 




i89I3 


I 


























~ 


































i0iI3 


m $ 

O 3" 

§ sr 


I 




















- 








































60iI3 


I 


























- 


































0UI3 


z 
























- 














- 






















TUI3 


T 
















































M 












ZIZ.I3 


I 
























- 




































£UI3 


681 


























^ 














b* 




- 




i 












6ZfrI3 


• 


S 


























•u 






















- 












IIH3 


in 
2. 
in si 

M ID 

§ 3 

ID "8 
u 

I- a 

— - 

en cr 
ID =< 

a £ 


£01 
























K. 


SI 








- 






- 






- 


to 












Zi*I3 


III 










- 














- 


NO 
ON 


w 






- 






- 








00 












fLPIO 


£E 
























- 
























N 












£8frI3 


Z 
















































to 












£69X3 


Z 
















































to 












W913 


► 


























■fe. 


































S69I3 


£ 
























- 


to 


































96913 


*• 


























■b 


































i69I3 


£ 


























UJ 


































86913 


I 


























- 


































66913 


I 


























{= 


































00U3 


i68 


- 


« 


- 





- 


- 


oc 


- 


'JJ 


-J 


- 


3 








- 


- 


A 


-J 


i. 





Ul 


H 


5 


to 


- 


- 


M 


O 


■U 


r«!J»x 



144 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



compressa Zone). 

The oldest fauna is represented by a small 
assemblage, including Appalachignathus delicatulus, 
Protopanderodus nogamii, IPeriodon aculeatus, 
Amelia sp., Erraticodon sp., and Stiptognathus sp. A 
from a single sample C1463, which was obtained from 
several small limestone clasts within the Fairbridge 
Volcanics at a stratigraphic level some 120 m below 
the Wahringa Limestone Member (Percival et al. 
1999). This fauna is comparable with that recently 
described from allochthonous limestones of Da3 age 
in the Oakdale Formation of the Bell River valley 
(Zhen and Percival in press), situated approximately 
23 km southeast of the "Wahringa" area. Of the six 
species recognised in sample CI 463, four also occur 
in the Bell River valley fauna. Stiptognathus sp. A is 
rare, but the other three species (Appalachignathus 
delicatulus, Protopanderodus nogamii, and 
Erraticodon sp.) dominate in all five samples from the 
Oakdale Formation (Zhen and Percival in press). On 
this basis, the age of sample C 1463 can now be revised 
downwards to the E. suecicus Zone from the 
previously-interpreted level near the Mid/Late 
Ordovician boundary (Zhen et al. 2001). 

In its type section, the Wahringa Limestone 
Member consists of three laterally continuous outcrops 
separated by two intervals of poor or negligible 
exposure (Fig. 2). Initially, one spot sample was 
collected from each of these three major outcrops, 
representing the lower, middle, and upper beds of the 
unit. Subsequent more intensive collecting during 
section measuring produced 17 samples that yielded 
conodonts. Although most of these samples have low 
yields and diversity, three conodont assemblages 
(herein referred to as A, B, and C from oldest to 
youngest) can be distinguished. 

Assemblage A was recovered from sample 
C 1 652 at the base of the Wahringa Limestone Member 
and spot sample C1450 within the lower part of this 
unit (essentially an equivalent stratigraphic level to 
C1652) in the type section. The fauna consists of 15 
species including Acodus sp., Ansella nevadensis, 
Ansella biserrata, Belodina sp. B, Dapsilodus 
variabilis, Drepanoistodus sp., Erraticodon balticusl, 
Oistodusl sp. cf. venustus, Panderodus gracilis, 
Periodon aculeatus, Phragmodus flexuosus, 
Protopanderodus nogamii, Protopanderodus 
varicostatus, Pygodus serra, and Pygodus 
protoanserinus. Most of these species are widely 
distributed and relatively long ranging, but the two 
species of Pygodus are important biostratigraphically. 
Pygodus protoanserinus has a range from the upper 
E. robustus Subzone to the E. lindstroemi Subzone 
(of the upper Pygodus serra Zone). One specimen (see 
Fig. 9K, L) referrable to the middle form of the Pa 



element of Pygodus serra (Zhang 1998a) was also 
recovered in the sample CI 65 2. Co-occurrence of P. 
serra and P. protoanserinus places the base of the 
Wahringa Limestone Member precisely within the 
upper E. robustus Subzone (upper Da3 to lowest Da4) 
of the P. serra Zone. 

Closest correlations are with successions in 
China. In the top Guniutan Formation (upper P. serra 
Zone) of Hunan Province, Zhang (1998b) recorded 
the co-occurrence of Pygodus protoanserinus with 
Erraticodon balticusl, Protopanderodus varicostatus, 
and Periodon aculeatus. Pygodus serra, Periodon 
aculeatus, Protopanderodus varicostatus, 
Protopanderodus nogamii, and Panderodus gracilis 
also occur in the lower part (P. serra Zone) of the 
Pingliang Formation of the Ordos Basin (An and Zheng 
1990). 

Only five samples from the middle section 
of the Wahringa Limestone Member have yielded 
conodonts. These faunas are of very low diversity and 
productivity, and are referred to herein as Assemblage 
B. They include Ansella nevadensis, Ansella sp., 
Belodina monitorensis, Panderodus gracilis, and 
Periodon aculeatus. Of these, only B. monitorensis is 
significant for age determination, occurring widely 
within the late Darriwilian to Gisbornian interval 
(Sweet in Ziegler 1981). Assemblage B is likely very 
close to the Mid/Late Ordovician boundary, probably 
within the Cahabagnathus sweeti Zone, although the 
precise recognition of the boundary within the middle 
Wahringa Limestone Member is not determinable on 
current evidence. 

The upper part of the type section of the 
Wahringa Limestone Member is relatively more 
productive, with 12 samples yielding an assemblage 
(designated as Assemblage C) of 15 species including 
Acodus sp., Ansella nevadensis, Belodina compressa, 
Belodina monitorensis, Besselodus sp., Dapsilodus 
variabilis, Dapsilodus viruensis, Drepanoistodus sp., 
Oistodusl sp. cf. venustus, Panderodus gracilis, 
Periodon aculeatus, Protopanderodus cooperi, 
Protopanderodus varicostatus, Protopanderodus 
liripipus and Stiptognathus sp. B. As a zonal index 
species of late Gisbornian equivalents in the North 
American Midcontinent zonal scheme, the occurrence 
of Belodina compressa in the upper part of the 
Wahringa Limestone Member indicates that top of this 
unit may be as young as Gi2. This species also occurs 
in six samples from limestone pods within the 
Fairbridge Volcanics above the Wahringa Limestone 
Member. The presence of two elements confidently 
identified as this species in limestone pods in the 
Fairbridge Volcanics slightly below the base of the 
Wahringa Limestone Member (samples C1487 and 
C1488) cannot be explained at present, as this 



Proc. Linn. Soc. N.S.W., 125, 2004 



145 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 




Figure 4. A-G, Acodus sp.; A, B, P element, MMMC2639, C1450, A, inner lateral view, B, outer lateral 
view; C, D, P element, MMMC2640, C1652, C, outer lateral view, D, anterior view; E-G, Sa element, 
MMMC2641, C1680, lateral views. H-P, Ansella nevadensis (Ethington and Schumacher 1969); H-J, Pa 
element, MMMC2642, C1450, H, inner lateral view, I, outer lateral view, J, showing surface striation; K, 
L, Pb element, MMMC2643, C1450, K, outer lateral view, L, inner lateral view; M, N, Sa element, 
MMMC2644, C1683, lateral views; O, P, Sc element, MMMC2645, C1450, 0, outer lateral view, P, inner 
lateral view. Q, Ansella biserrata Lehnert and Bergstrom in Lehnert et al. 1999; Pa element, MMMC2646, 
C1652, outer lateral view. R, S, Ansella sp.; R, Pb element, MMMC2647, C1486, inner lateral view; S, Pb 
element, MMMC2648, C1672, outer lateral view. T, Appalachignathus delicatulus Bergstrom et al. 1974; 
Pb element, MMMC2649, C1463, inner lateral view. Unless otherwise indicated scale bars are 100 pjn. 



146 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



contradicts the species succession in all known global 
occurrences. 



LOCALITIES AND SAMPLES 

Details of the localities and measured section 
are shown in Figures 1 and 2, and summarised in the 
Appendix. Distribution of conodont species is 
presented in Figure 3. All illustrations of conodont 
elements are presented as SEM photomicrographs 
(Figs 4-9). Figured specimens bearing the prefix 
MMMC ("Mining Museum microfossil catalogue") 
are deposited in the collections of the Geological 
Survey of New South Wales, Sydney. Individual 
samples are referred to by the prefix "C". Although 
all species recorded are documented by illustration, 
only those where adequate material was recovered, or 
which are of biostratigraphic significance, are 
described. Unless otherwise mentioned, all specimens 
are from the Wahringa Limestone Member. 



SYSTEMATIC PALAEONTOLOGY 

Class CONODONTATA Pander 1856 

Genus ANSELLA Fahraeus and Hunter 1985a 

Type species 

Belodella jemtlandica Lofgren 1978. 

Amelia nevadensis (Ethington and Schumacher 
1969) 
Fig. 4H-P 
Synonymy 
Roundya sp. Sweet and Bergstrom 1962, p. 1244, 

1245, text-fig. 5. 
New Genus A Ethington and Schumacher 1969, p. 

478, 479, pi. 68, fig. 12, text-fig. 4J. 
Oepikodus copenhagenensis Ethington and 

Schumacher 1969, p. 465, pl.68, figs 5, 9, 

text-fig. 4L. 
Oistodus nevadensis Ethington and Schumacher 

1969, p. 467, 468, pi. 68, figs 1-4, text-fig. 

5C; Tipnis et al. 1978, pi. 6, fig. 7. 
Belodella nevadensis (Ethington and Schumacher); 

Bergstrom 1978, pi. 79, figs 9, 10; Bauer 

1987, text-fig. 5D. 
Ansella nevadensis (Ethington and Schumacher); 

Fahraeus and Hunter 1985a, p. 1175, 1176, 

pi. 1, figs 7, 10, pi. 2, figs 11a, b, 13a, b, 

14, text-fig. 2A-C; Bergstrom 1990, pi. 1, 

figs 11-14; McCracken 1991, p. 47-49, pi. 

3, figs 3, 4, 8, 9, 13, 14, 19-31 (cum syn.); 



?Bauer 1990, pi. 1, fig. 1; ?Bauer 1994, fig. 
3.4, 3.5. 

Material 

Ten specimens (1 Pa, 4 Pb, 4 Sa, 1 Sc). 

Description 

The P elements are characterised by a prominent 
median costa on each side, and display a sharply inner 
laterally curved anterior margin. The Pa element has a 
row of denticles along the posterior edge (Fig. 4H-J). 
The denticle next to the cusp is the largest, and the 
others become gradually smaller towards the base. A 
sharp costa on each side extends from the tip of the 
cusp and disappears a short distance away from the 
basal margin. The Pb element has a sharp posterior 
margin without any denticles, a weaker and broader 
costa on the inner lateral side, and a sharp, strong costa 
on the outer lateral side (Fig. 4K, L). The Sa element 
is symmetrical with a row of closely spaced small 
denticles along the posterior margin, and bears a sharp 
antero-lateral costa on each side (Fig. 4M, N). The 
asymmetrical Sc element has an inner laterally curved 
anterior margin, and a row of small closely spaced 
denticles along the posterior margin (Fig. 40, P). 
Specimens are ornamented with fine striation. 

Discussion 

Originally proposed as the form species Oepikodus 
copenhagenensis Ethington and Schumacher 1969 and 
New Genus A Ethington and Schumacher 1969 (found 
in association with Oistodus nevadensis in the 
Copenhagen Formation of Nevada), these elements 
were considered as part of the species apparatus of A. 
nevadensis by Fahraeus and Hunter (1985a), and are 
herein assigned to the Pa and Sb positions. Fahraeus 
and Hunter (1985a) also illustrated a symmetrical Sa 
element (Fahraeus and Hunter 1985a, pi. 2, fig. 14) 
from the Cobbs Arm Limestone of Newfoundland. 
Specimens from the Wahringa Limestone Member 
permit differentiation of denticulate Pa and 
adenticulate Pb elements. The latter has not been 
recognised previously, but its assignment to the Pb 
position is consistent with the apparatus composition 
of comparable species such as A. jemtlandica and A. 
crassa Bauer 1994, from central New South Wales 
(Zhen and Percival in press). 

Ansella biserrata Lehnert and Bergstrom in Lehnert 
et al. 1999 
Fig. 4Q 
Synonymy 

Ansella biserrata Lehnert and Bergstrom in Lehnert 
et al. 1999, p. 210, 212, pi. 1, figs 4, 7, pi. 
3, figs 1-3, 5 (cum syn.). 



Proc. Linn. Soc. N.S.W., 125, 2004 



147 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 



Material 

One specimen (Pa). 

Discussion 

Lehnert and Bergstrom in Lehnert et al. (1999) 
recognised a quadrimembrate apparatus for A. 
biserrata including biserrate, planoconvex, 
oistodiform, and triangular elements. We refer the 
biserrate and planoconvex elements to the Pa and Pb 
positions respectively, whereas the triangular element 
is regarded as taking either the Sa (symmetrical) or Sb 
(asymmetrical) position. The sole specimen from the 
lower Wahringa Limestone Member (CI 652), with 
smooth lateral faces and fine denticles along its anterior 
and posterior margins, strongly resembles the holotype 
(a biserrate element) of A. biserrata from the basal 
Lindero Formation (Pygodus serra and P. anserinus 
zones) of west central Argentina (Lehnert et al. 1999). 

Amelia sp. 
Fig. 4R, S 

Synonymy 

Serraculodusl sp. Zhen and Webby 1995, p. 286, 

only pi. 5, figs 1, 3, 4. 
Amelia sp. Zhen et al. 2003a, p. 38, fig. 4A, 4B. 

Material 

Three specimens (Pb). 

Discussion 

These specimens are similar to the Pb element of A. 
nevadensis, but they lack the prominent lateral median 
costae of that species. They are identical with some 
specimens from the Fossil Hill Limestone of Eastonian 
age at Cliefden Caves previously assigned to 
Serraculodusl sp. (Zhen and Webby 1995). They can 
be distinguished from the Pb elements of both A. 
jemtlandica (Lofgren 1978) and A. crassa Bauer 1994 
in lacking the posteriorly more expanded base 
displayed in the latter two species (Zhen and Percival 
in press). 

Genus BELODINA Ethington 1959 

Type species 

Belodus compressus Branson and Mehl 1933. 

Belodina compressa (Branson and Mehl 1933) 
Fig. 5A-I 
Synonymy 
Belodus compressus Branson and Mehl 1933, p. 

114. pi. 9, figs 15, 16. 
Belodus grandis Stauffer 1935, p. 603-604, pi. 72, 
figs 46, 47, 49, 53, 54, 57. 



Belodus wykoffensis Stauffer 1935, p. 604, pi. 72, 
figs 51, 52, 55, 58, 59. 

Oistodus fornicalus Stauffer 1935, p. 610, pi. 75, 
figs 3-6. 

Belodina dispansa (Glenister); Schopf 1966, p. 43, 
pl. 1, fig- 7. 

Belodina compressa (Branson and Mehl); Bergstrom 
and Sweet 1966, p. 321-315, pl. 31, figs 
12-19; Webers 1966, p. 24, pl. 1, figs 2, 6, 
7, 13, 15; Sweet in Ziegler 1981, p. 65-69, 
Belodina - plate 2, figs 1-4; An et al. 1983, 
only pl. 25, figs 13, 14; Moskalenko 1983, 
fig. 3Q-S; Leslie 1997, p. 921-926, figs 
2.1-2.20, 3.1-3.4 (cum syn.). 

Belodina confluens Sweet; Percival et al. 1999, p. 
13, fig. 8.21. 

Material 

255 specimens, including eobelodiniform, 
compressiform, grandiform and dispansiform 
elements, mainly from the upper part of the Wahringa 
Limestone Member; some specimens from 
allochthonous limestones in the Fairbridge Volcanics 
above the Wahringa Limestone Member, and two 
elements from limestone pods (C1487, C1488) in the 
Fairbridge Volcanics which apparently underlie the 
Wahringa Limestone Member. 

Discussion 

Of the known species of Belodina, three including B. 
compressa, B. confluens Sweet 1979, and B. 
monitorensis Ethington and Schumacher 1969, are 
morphologically very similar to each other, reflecting 
their close phylogenetic relationship. Well- 
documented successions in the U.S.A. (Sweet 1979) 
show that the oldest species, B. monitorensis, preceded 
B. compressa which was succeeded by B. confluens. 
Sweet (in Ziegler 1981, p. 65) revised all three species 
as consisting of trimembrate apparatuses, and 
emphasised that the type species of the genus, B. 
compressa, was characterised by having a distinct 
flattening (in lateral view) of the anterior margin near 
the antero-basal corner. This feature is more prominent 
in the compressiform element, as shown by the types 
(Branson and Mehl 1933) and also the specimens 
figured by Webers (1966); also see Sweet in Ziegler 
(1981). Both B. confluens and B. compressa are 
commonly found in association with a more slender, 
rastrate element bearing smaller denticles. Bergstrom 
(1990) suggested that these dispansiform elements 
might represent juveniles of the rastrate elements. 
Many other workers included these dispansiform 
elements in a separate species (dispansa) assigned 
either to Pseudobelodina (Sweet in Ziegler 1981, 
Nowlan and McCracken in Nowlan et al. 1988, 



148 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 




Figure 5. A-I, Belodina compressa (Branson and Mehl 1933); A, inner lateral view, B, outer lateral view, 
grandiform element, MMMC2650, C1472; C, dispansiform element, MMMC2651, C1429, inner lateral 
view; D, compressiform element, MMMC2652, C1472, inner lateral view; E, compressiform element, 
MMMC2653, C1458, outer lateral view; F, compressiform element, MMMC2654, C1683, outer lateral 
view; G, dispansiform element, MMMC2655, C1429, outer lateral view; H, inner lateral view, I, outer 
lateral view, eobelodiniform element, MMMC2656, C1683. J-N, Belodina monitorensis Ethington and 
Schumacher 1969; J, outer lateral view, K, inner lateral view, grandiform element, MMMC2657, C1687; 
L, inner lateral view, M, outer lateral view, compressiform element, MMMC2658, C1456; N, 
eobelodiniform element, MMMC2659, C1456, outer lateral view. O, P, Belodina sp. B; O, eobelodiniform 
element, MMMC2660, C1450, inner lateral view; P, eobelodiniform element, MMMC2661, C1652, outer 
lateral view. Q, Belodina sp. A; grandiform element, MMMC2662, C1429, outer lateral view. R-T, 
Besselodus sp.; R, S, Sa element, MMMC2663, C1676, lateral views; T, M element, MMMC2664, C1675, 
anterior view. Scale bars are 100 urn. 



Proc. Linn. Soc. N.S.W., 125, 2004 



149 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 



McCracken and Nowlan 1989, Trotter and Webby 
1995, Leslie 1997, McCracken 2000) or to Belodina 
(Schopf 1966, Barnes 1977, Nowlan and Barnes 1981, 
Sansom et al. 1995). In comparing the apparatus 
architecture of Panderodus, Sansom et al. (1995) 
suggested that these slender dispansiform elements, 
which were previously included in Belodina dispansa 
(Glenister 1957) and Belodina area Sweet 1979, might 
belong to the species apparatus of co-occurring B, 
confluens. Based on twelve well preserved, fused 
clusters of B. compressa recovered from the Plattin 
Limestone of Missouri and Iowa, Leslie (1997) 
demonstrated that B. compressa consisted of a 
quadrimembrate apparatus including the M 
(eobelodiniform), SI (compressiform), S2 
(grandiform) and S2 (dispansiform) elements. He 
further suggested that the dispansiform element - 
although superficially similar in morphology to 
Pseudobelodina dispansa - was apparently not 
conspecific. Leslie also rejected the possibility that 
such elements represented the juveniles of 
compressiform and grandiform elements in 
consideration of the range of sizes and growth series 
preserved in the dispansiform elements. 

The material of B. compressa and B. 
confluens from central New South Wales shows 
recognisable differences between the two species. 
Compressiform elements of B. compressa display in 
lateral view a straight segment of the anterior margin 
near the antero-basal corner. In comparison, the 
anterior margin of the compressiform element of B. 
confluens is regularly curved near the antero-basal 
corner. Hence specimens of B. compressa previously 
reported from the Fork Lagoons Beds of central 
Queensland (Palmieri 1978), and from the Trelawney 
Beds of the New England Fold Belt (Philip 1966) were 
subsequently reassigned to B. confluens (Zhen and 
Webby 1995). Specimens from the Wahringa 
Limestone Member and various limestone pods within 
the Fairbridge Volcanics are the first confirmed records 
of B. compressa from eastern Australia. 

Belodina monitorensis Ethington and Schumacher 
1969 
Fig. 5J-N 
Synonymy 

Belodina monitorensis monitorensis Ethington and 
Schumacher 1969, p. 456, pi. 67, figs 3, 5, 
8, 9, text-fig. 5D. 
Belodina monitorensis marginata Ethington and 

Schumacher 1969, p. 456, pi. 67, figs 1, 2, 
4. 6, text-fig. 5E. 
Eobelodina occidentalis Ethington and Schumacher 
1969, p. 456, pi. 67, figs 16, 20, text-fig. 
5H. 



Belodina monitorensis Ethington and Schumacher 
1969, p. 455, 456; Sweet in Ziegler 1981, 
p. 79-81, Belodina - plate 1, figs 10, 1 1; 
Belodina - plate 2, figs 5-7; Bauer 1987, p. 
12, pi. 1, figs 10, 13, 14; Bauer 1990, pi. 1, 
fig. 9; Bauer 1994, fig. 3.16, 3.17, 3.20, 
3.21. 

Material 

17 specimens including eobelodiniform, 
compressiform and grandiform elements. 

Discussion 

Belodina monitorensis was originally defined as having 
prominent antero-lateral costae on either side of the 
grandiform element and generally four or five denticles 
on both grandiform and compressiform elements. A 
similar antero-lateral costa is also commonly found in 
the grandiform elements of B. compressa (Fig. 5B; 
also see Leslie 1997, fig. 2.3), and in the grandiform 
elements of B. confluens (McCracken 1987, pi. 1, fig. 
1; Zhen and Webby 1995, pi. 1, figs 17, 20; Zhen et 
al. 1999, fig. 5.8). Therefore, this character appears to 
be unreliable in characterising B. monitorensis. 
Although B. confluens and B. compressa typically have 
a greater number of denticles (five to nine), it seems 
rather arbitrary to split B. monitorensis (typically four 
or five denticles) from B. confluens based solely on 
the former having fewer denticles. 

Though the species status of B. monitorensis 
is uncertain in our view, stratigraphically it occurs 
much earlier than typical B. confluens. In the type 
section of the Wahringa Limestone Member, B. 
monitorensis occurs lower than B. compressa, but it is 
also found in association with the latter species in 
several samples in the upper part of the type section. 
The Wahringa Limestone Member specimens are 
comparable with the type material of B. monitorensis 
in having only three or four denticles, and in having 
an antero-lateral costa on the furrowed side of the 
grandiform element (Fig. 5J), also shown by the 
holotype (Ethington and Schumacher 1969, pi. 67, fig. 
5). Therefore, the species is tentatively retained here 
pending further detailed studies on B. monitorensis and 
other related species. 

Belodina sp. A 
Fig. 5Q 
Material 

One specimen from sample C1429 (upper beds of the 
Wahringa Limestone Member at the southwestern 
extremity of its outcrop). 

Discussion 

This compressiform element has a squat cusp and two 



150 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



short and stout denticles along the posterior margin. 
The specimen is not as strongly compressed laterally 
as other known species of Belodina. 

Belodina sp. B 
Fig. 50, P 
Material 

Three eobelodiniform specimens. 

Discussion 

From its association with Pygodus protoanserinus in 
two samples (C1450 and C1652) from the lower part 
of the Wahringa Limestone Member this species has a 
late Darriwilian age (upper Da3 to lowest Da4, upper 
P. serra Zone), making it one of the earliest 
representatives of the genus Belodina. With a less 
extended heel it shows some morphological 
resemblance to the eobelodiniform element of 
Belodina beiyanhuaensis Qiu in Lin, Qiu and Xu 1984, 
but no rastrate elements have been recovered to 
confirm such an assignment. 

Genus ERRATICODON Dzik 1978 

Type species 

Erraticodon balticus Dzik 1978. 

Erraticodon balticus? Dzik 1978 
Fig. 6P, Q 

Synonymy 

Erraticodon balticus Dzik 1978, p. 66, pi. 15, figs 1- 
3, 5, 6, text-fig. 6; ?Stouge 1984, p. 84, pi. 
17, figs 9-19; Watson 1988, p. 113, pi. 5, 
figs 2-10, pi. 8, figs 1, 2, 5, 6, 8-13 (cum 
syn.); Dzik 1991, p. 299, fig. 12A; Ding et 
al. in Wang 1993, p. 176, pi. 37, only figs 
19-28, non fig. 18; ?Pohler 1994, pi. 3, figs 
3-5; Lehnert 1995, p. 87, pi. 10, figs 13, 16, 
pi. 12, figs 3-5; ?Zhang 1998b, p. 71, pi. 9, 
figs 6-13; ?Albanesi in Albanesi et al. 
1998, p. 176, pi. 4, figs 16-18; ?Johnston 
and Barnes 2000, p. 19, pi. 4, figs 18, 20, 
23, 24, 29; Zhao et al. 2000, p. 203, pi. 36, 
figs 1-16; ?Pyle and Barnes 2002, p. Ill, 
pl.20, figs 8, 9. 

Material 

One specimen (M). 

Discussion 

Dzik (1978) originally defined the species as consisting 
of a seximembrate apparatus, but later (Dzik 1991) 
determined a septimembrate apparatus with digyrate 
Pa and Pb elements as typical of the species (Zhen et 
al. 2003b). Erraticodon balticus is characterised by 



having an accentuated denticle on the posterior process 
of the Sa, Sb and Sc elements (Dzik 1991, fig. 12A). 
The specimen from the Wahringa Limestone Member 
is broadly comparable with the M element of the 
illustrated type material (Dzik 1978, pi. 15, fig. 5), 
except that the latter has a reclined cusp; as our 
specimen has an erect cusp, it is only questionably 
referred to this species. 

Specimens ascribed to Erraticodon balticus 
from the Guniutan Formation of South China (Zhang 
1998b), the San Juan Formation of the Precordillera 
in Argentina (Albanesi in Albanesi et al. 1998), the 
Ospika Formation of British Columbia (Pyle and 
Barnes 2001), and the Cow Head Group of western 
Newfoundland (Johnston and Barnes 2000), all 
apparently lack the distinctive larger denticle on the 
posterior process of the S elements, and therefore 
should only be doubtfully assigned to the species. 

Erraticodon sp. 
Fig. 60 
Material 

One specimen (Sa) from sample C1463, a limestone 
pod in the Fairbridge Volcanics stratigraphically below 
the Wahringa Limestone Member. 

Discussion 

This alate element is identical with the Sa element of a 
new species of Erraticodon under description from 
allochthonous limestones within the Oakdale 
Formation of central New South Wales (Zhen and 
Percival in press). It has a prominent cusp with a 
flange-like costa on each side, which extends basally 
to define the upper margin of the lateral processes. 
The lateral processes bear four widely spaced, peg- 
like denticles. Comparison with other species of 
Erraticodon are discussed elsewhere (Zhen and 
Percival in press). 

Genus PEPJODON Hadding 1913 

Type species 

Periodon aculeatus Hadding 1913. 

Periodon aculeatus Hadding 1913 
Figs 6R, S, 7A-K 

Synonymy 

Periodon aculeatus Hadding 1913, p. 33, pi. 1, fig. 
14; Lindstrom 1955b, p. 110, pi. 22, figs 
10, 11, 14-16, 35; Lofgren 1978, p. 74, pi. 
10, fig. 1; pi. 11, figs 12-26, Fig. 29 (cum 
syn.); Sweet in Ziegler 1981, p. 237, 
Periodon - plate 1, figs 1-6; Nowlan 1981, 
pi. 4, figs 1-9; An 1987, p. 167, pi. 24, figs 
7-17; Bergstrom 1990, pi. 1, figs 15, 16, pi. 



Proc. Linn. Soc. N.S.W., 125, 2004 



151 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 




Figure 6. A-D, Dapsilodus variabilis (Webers 1966); A, B, symmetrical distacodontiform element, 
MMMC2665, C1675, lateral views; C, symmetrical distacodontiform element, MMMC2666, C1450, lateral 
view; D, acodontiform element, MMMC2667, C1652, outer lateral view. E-J, Dapsilodus viruensis 
(Fahraeus 1966). E, F, Sa element, MMMC2668, C1675, lateral views; G, outer lateral view, H, inner 
lateral view, Sb element, MMMC2669, C1675; I, outer lateral view, J, inner lateral view, Sc element, 
MMMC2670, C1675. K-M, Drepanoistodus sp.; K, outer lateral view, L, inner lateral view, Sc element, 
MMMC2671, C1652; M, P element, MMMC2672, C1450, inner lateral view. N, Oistodusl sp. cf. venustus 
Stauffer 1935; anterior view, MMMC2673, C1450. O, Erraticodon sp.; Sa element, MMMC2674, C1463, 
postero-lateral view. P, Q, Erraticodon balticusl Dzik 1978; M element, MMMC2675, C1450, P, posterior 
view, Q, anterior view. R, S, Periodon aculeatus Hadding 1913; Pb element, MMMC2676, C1450, R, 
inner lateral view, S, outer lateral view. Scale bars are 100 urn. 



152 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



2, fig. 15; An and Zheng 1990, pi. 12, figs 
12-17; McCracken 1991, p. 50, pi. 1, figs 
13, 20, 22, 25-28, pi. 2, figs 24-27, 31, 34, 
35 (cum syn.); Pohler 1994, pi. 4, figs 30- 
32; Dzik 1994, p. Ill, pi. 24, figs 10-13, 
text-fig. 31b; Lehnert 1995, p. 110, pi. 10; 
fig. 2, pi. 11, figs 10, 11, pi. 13, figs 9, 11, 
12, pi. 16, figs 8, 9, 11-13; Armstrong 
1997, p. 774, pi. 2, figs 13-21; text-fig. 3; 
Albanesi in Albanesi et al. 1998, p. 170, pi. 
15, figs 16-17, pi. 16, figs 19, 20 (cum 
syn.); Zhang 1998b, p. 80, 81, pi. 14, figs 
1-8 (cum syn.); Johnston and Barnes 2000, 
p. 32-35, pi. 13, figs 12, 13, 17, 18, 20-31, 
pi. 14, figs 1-7, text-figs 4, 5 (cum syn.); 
Rasmussen 2001, p. 110, pi. 13, figs 8-11 
(cum syn.); Pyle and Barnes 2002, p. 107, 
pi. 21, figs 7-9. 

Material 

97 specimens. 

Description 

Both Pa and Pb elements are angulate with a prominent 
cusp which is laterally compressed with a median costa 
on each side. The Pb element differs from the Pa 
element in having a twisted posterior process and a 
strongly inner laterally curved and downwardly 
extended anterior process (Fig. 6R, S). The M element 
is makellate with an adenticulate outer lateral process, 
and with 4-6 closely spaced denticles along the inner 
lateral margin. The alate Sa element has a long 
posterior process bearing closely spaced denticles. The 
sixth denticle away from the cusp is much larger and 
more robust (Fig. 7E, F). The lateral process on each 
side is blade-like, bearing small closely spaced 
rudimentary denticles along the edge. The basal cavity 
is shallow with a recessive basal margin zone. The Sb 
element is tertiopedate and asymmetrical, and bears a 
long denticulate posterior process with closely spaced, 
strongly reclined denticles, a long inner lateral process 
with more than seven small confluent denticles, and a 
short outer lateral process with only two small denticles 
(Fig. 7H, G). The Sc element is bipennate with a long, 
denticulate posterior process, bearing closely spaced, 
strongly reclined denticles, and with an inner laterally 
curved anterior process bearing small confluent 
denticles (Fig. 7I-K). 



alate Sa, tertiopedate Sb and bipennate Sc, elements. 
Albanesi (in Albanesi et al. 1998, text-fig 31, pi. 9, 
fig. 10) suggested a septimembrate apparatus for the 
species by recognizing a lozognathiform Sd element, 
which bears a long denticulate, twisted posterior 
process, a short, denticulate outer lateral process and 
an inner laterally curved, sharp, blade-like anterior 
costa. Rasmussen (2001, pi. 13, fig.l 1) also recognised 
a modified tertiopedate Sd element, and described it 
as characterised by a multidenticulate, twisted, 
posterior process and weakly denticulated anterior 
process, and process-like extension of the outer-lateral 
costa or carina, but only a poorly preserved specimen 
was illustrated. In the Wahringa collections no Sd 
elements have been recognised. 

Lofgren (1978, p. 75) suggested that the 
number of small denticles between the cusp and the 
biggest denticle increases from a mean of 4.7 to 5.6 in 
successively younger samples. Specimens from the 
Wahringa Limestone Member may therefore represent 
more advanced forms of the species within its 
stratigraphic range, as shown by the M elements, which 
are strongly geniculate with a sinuous basal margin 
and bear 4-6 denticles (mean 5.5) along the inner lateral 
margin. 

Genus PHRAGMODUS Branson and Mehl 1933 

Type species 

Phragmodus primus Branson and Mehl 1933. 

Phragmodus flexuosus Moskalenko 1973 
Fig. 7L, M 

Synonymy 

Phragmodus sp. Moskalenko 1972, p. 48-50, text- 
fig. 1, table 2. 

Phragmodus flexuosus Moskalenko 1973, p. 73, 74, 
pi. 11, figs 4-6; Sweet in Ziegler 1981, p. 
255-258, Phragmodus - plate 2, figs 1-6 
(cum syn.); Bauer 1987, p. 24, 25, pi. 3, 
figs 10, 14, 15, 17, 18, 20, 24, text-fig. 8; 
Ethington and Clark 1982, p. 79-82, pi. 9, 
figs 2-7 (cum syn.); Bauer 1994, p. 367, 
368, fig. 5.25, 5.26, 5.28, 5.30-5.33; 
Percival et al. 1999, fig. 8.15. 

Material 

One specimen (Sa). 



Discussion 

Following the Treatise definition of the genus (Clark 
et al. 1981, p. W128), Sweet (1988) proposed a 
seximembrate apparatus for P. aculeatus, consisting 
of angulate Pa and Pb, makellate M, and ramiform 



Discussion 

This specimen is alate with a suberect cusp, a long 
denticulated posterior process, and with a flange-like 
costa on each lateral side. The straight posterior process 
supports more than seven widely spaced denticles, 



Proc. Linn. Soc. N.S.W., 125, 2004 



153 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 




Figure 7. A-K, Periodon aculeatus Hadding 1913; A, outer lateral view, B, inner lateral view, Pa element, 
MMMC2677, C1450; C, posterior view, D, anterior view, M element, MMMC2678, C1450; E, F, Sa 
element, MMMC2679, C1450, lateral views; G, outer lateral view, H, inner lateral view, Sb element, 
MMMC2680, C1450; I, outer lateral view, J, inner lateral view, Sc element, MMMC2681, C1450; K, Sc 
element, MMMC2682, C1652, outer lateral view. L, M, Phragmodus flexuosus Moskalenko 1973; Sa 
element, MMMC2683, C1450, lateral views. N-U, Panderodus gracilis (Branson and Mehl 1933); N, 
posterior view, O, inner lateral view, P, outer lateral view, graciliform element, MMMC2684, C1450; Q, 
posterior view, R, basal view, S, lateral view, aequaliform element, MMMC2685, C1458; T, falciform 
element, MMMC2686, C1697, outer lateral view; U, falciform element, MMMC2687, C1458, inner lateral 
view. Scale bars are 100 urn. 



which are reclined, similar in size, with V- or U-shaped 
spaces between. It is comparable to the type material 
from Siberia, except that the Wahringa specimen 
exhibits a rather straight posterior process with more 



or less equal-sized denticles, and a few small, 
rudimentary denticles on the lower edge of the lateral 
processes. Although Moskalenko (1972) initially 
recognised nine morphotypes for the species, she later 



154 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



(Moskalenko 1973) formally described the species as 
a form species. Based on a detailed revision, Ethington 
and Clark (1982) redefined P. flexuosus as consisting 
of a seximembrate apparatus. However, the illustrated 
S elements from Utah display more pronounced 
undulations and twisting of the posterior process and 
size variation of the denticles on the posterior process 
(Ethington and Clark 1982, pi. 9, figs 3, 6) than the 
type material from Siberia (Moskalenko 1973). 

Genus PROTOPANDERODUS Lindstrom 1971 

Type species 

Acontiodus rectus Lindstrom 1955a. 

Protopanderodus cooperi (Sweet and Bergstrom 

1962) 

Fig. 8A-E 

Synonymy 

Scandodus rectus Lindstrom 1955a, p. 593, only pi. 

4, figs 22, 23. 
Acontiodus cooperi Sweet and Bergstrom 1962, p. 

1221, pi. 168, figs 2, 3, text-fig. 1G. 
Scandodus sp. Sweet and Bergstrom 1962, p. 1246, 

pi. 168, figs 13, 16. 
Protopanderodus cooperi (Sweet and Bergstrom); 
Zhang 1998b, p. 81, 82, pi. 14, figs 13-17 
(cum syn.). 



than one costa on each lateral side. 

Zhang (1998b) provided a comprehensive 
synonymy list, and illustrated what she recognised as 
P, M, Sa and other undifferentiated S elements; 
however, Zhang provided neither diagnosis nor 
descriptions of the constituent elements of the species 
apparatus of P. cooperi. This species was originally 
proposed as a form species from the Ferry Formation 
of Alabama. The holotype (Sweet and Bergstrom 1962, 
pi. 168, figs 2, 3) is slightly asymmetrical, defined here 
as taking the Sb position. Zhang (1998b) included the 
form species Scandodus sp. Sweet and Bergstrom 1962 
in the P position of P. cooperi. Based on their 
illustrations and brief discussion, the P element is 
inferred to be a scandodiform element with broad costa 
on the inner lateral face and with a smooth outer lateral 
face. Zhang (1998b) also included the holotype of 
Scandodus rectus Lindstrom 1955a as occupying the 
M position in P. cooperi. This scandodiform element 
is similar to the P element previously defined, except 
that only a broad carina is developed on the inner lateral 
face. For consistency, these two scandodiform 
elements are tentatively taken to represent the Pa and 
Pb positions in Protopanderodus. The symmetrical Sa 
of P. cooperi was illustrated from the Guniutan 
Formation of South China (Zhang 1998b, pi. 14, fig. 
13), and was also recovered from the Wahringa 
samples (Fig. 8A, B). 



Material 

Seven specimens (6 Sa, 1 Sb). 



Protopanderodus robustus (Hadding 1913) 
Fig. 8J-M 



Discussion 

This species is rare in the Wahringa collections. Two 
morphotypes are recognised as representing the Sa and 
Sb elements, all with sharp anterior and posterior 
margins, and a suberect cusp and one costa on each 
lateral side. The Sa element is symmetrical with a sharp 
median costa (Fig. 8A, B). The Sb element resembles 
the Sa, but is slightly asymmetrical with a more 
strongly developed costa on the inner side (Fig. 8C- 
E). No scandodiform P elements and no laterally 
compressed Sc elements, as characterising P. cooperi 
of previous authors, were recovered. Based on the 
original definition of the species given by Sweet and 
Bergstrom (1962) and more recent revision (Zhang 
1998b), elements of P. cooperi exhibit sharp anterior 
and posterior margins, a well developed anticusp-like 
extension at the antero-basal corner, deep antero-lateral 
recesses in the basal margin, and no more than one 
costa on each lateral face. Protopanderodus cooperi 
can be differentiated from P. rectus (Lindstrom) in 
having an anticusp-like extension at the anterobasal 
corner, »nd from P. varicostatus in displaying no more 



Synonymy 

Drepanodus robustus Hadding 1913, p. 31, pi. 1, 

fig.5. 
Protopanderodus robustus (Hadding); Lofgren 

1978, p. 94, 95, pi. 3, figs 32-35, text-fig. 

31G-J (cum syn.); An 1987, p. 173, pi. 11, 

figs 7-10 (cum syn.); McCracken 1989, p. 

20-22, pi. 1, figs 1-10, text-fig. 3E (cum 

syn.); Albanesi in Albanesi et al. 1998, p. 

129, 130, pi. 11, figs 17-20, text-fig. 14A 

(cum syn.). 

Material 

Two specimens (Sa, Sc). 

Discussion 

One specimen in the Wahringa collection (Fig. 8L, 
M) which has sharp anterior and posterior margins and 
is laterally compressed with a postero-lateral costa on 
each side, is regarded as representing the Sa element 
of Protopanderodus robustus. The other specimen with 
a single costa on the outer lateral face is referred to the 



Proc. Linn. Soc. N.S.W., 125, 2004 



155 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 




Figure 8. A-E, Protopanderodus cooperi (Sweet and Bergstrom 1962); A, B, Sa element, MMMC2688, 
C1683; lateral views; C, upper view, D, outer lateral view, E, inner lateral view, Sb element, MMMC2689, 
C1682. F-I, Protopanderodus nogamii (Lee 1975); F-H, Sa element, MMMC2690, C1450, lateral views; I, 
Sa element, MMMC2691, C1463, lateral view. J-M, Protopanderodus robustus (Hadding 1913); J, outer 
lateral view, K, inner lateral view, Sc element, MMMC2692, C1680; L, M, Sa element, MMMC2693, 
C1458, lateral views. N-X, Protopanderodus varicostatus (Sweet and Bergstrom 1962); N, Pa element, 
MMMC2694, C1652, inner lateral view; O, outer lateral view, P, inner lateral view, Pb element, 
MMMC2695, C1675; Q, R, Sd element, MMMC2696, C1675, lateral views; S, T, Sd element, MMMC2697, 
C1682, lateral views; U, outer lateral view, V, inner lateral view, Sb element, MMMC2698, C1675; W, 
inner lateral view, X, outer lateral view, Sc? element, MMMC2699, C1675. Y, Z, Protopanderodus liripipus 
Kennedy et al. 1979; Sa element, MMMC2700, C1458, lateral views. Scale bars are 100 urn. 



156 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



Sc element of the same species (Fig. 8J, K). Although 
Lindstrom (1971), and Johnston and Barnes (2000) 
maintained the original generic assignment of the form 
species Drepanodus robustus Hadding 1913, most 
other authors followed the revision of Lofgren (1978) 
who regarded it as a species of Protopanderodus. 
Lofgren (1978) recognised three morphotypes: 
symmetrical acontiodiform (an Sa element) with a 
postero-lateral costa on each side (Fig. 8L, M), 
asymmetrical acontiodiform (an Sc element) with a 
strong costa on the outer lateral face and a non costate 
inner lateral face (Fig. 8J, K), and scandodiform 
(interpreted here as a P element). Based on this 
multielement species definition, P. robustus is 
morphologically very close to P. cooperi. Specimens 
referable to D. robustus were also recorded from the 
Pratt Ferry Formation of Alabama, where the type 
material of P. cooperi was described (Sweet and 
Bergstrom 1962). Zhang (1998b) suggested that the 
holotype of D. robustus might be an element of an 
uncertain species of Protopanderodus, but she included 
all the material described from Sweden by Lofgren 
(1978) as P. robustus (Hadding) in her synonymy of 
P. cooperi. This implies that P. cooperi may be a junior 
synonym of P. robustus. Given that the base of the 
holotype of Drepanodus robustus is apparently broken 
(see also Lindstrom 1955b), it remains difficult to 
separate these two species. 

Protopanderodus varicostatus (Sweet and 

Bergstrom 1962) 

Fig. 8N-X 

Synonymy 

Scolopodus varicostatus Sweet and Bergstrom 1962, 
p. 1247, pi. 168, figs 4-9, text-fig. 1A, C, 
K. 

Scandodus unistriatus Sweet and Bergstrom 1962, 
p. 1245, pi. 168, fig. 12, text-fig. IE. 

Protopanderodus varicostatus (Sweet and 

Bergstrom); Dzik 1976, only text-fig. 16f, 
g; Fahraeus and Hunter 1985b, p. 183, text- 
fig. 2; Bauer 1987, p. 27, pi. 3, figs 19, 21- 
23; An 1987, p. 173, pi. 11, figs 2, 3; Dzik 
1994, p. 74, pi. 14, figs 1-5, text-fig. lib; 
Zhang 1998b, p. 83, 84, pi. 15, figs 14-19 
(cum syn.). 

Material 

Seven specimens from sample CI 675, and one 
specimen from C1652. 

Discussion 

Sweet and Bergstrom (1962) originally recognised 
three form-groups for the species, namely symmetrical, 
slightly asymmetrical, and markedly asymmetrical. 



Fahraeus and Hunter (1985b) proposed a 
quinquimembrate apparatus for this species, with 
elements referred to as groups A to E. Group A is a 
symmetrical multi-costate element with two costae on 
each lateral face. Group B is an asymmetrical tri- 
costate element (= the markedly asymmetrical form 
group of Sweet and Bergstrom 1962) with two costae 
on the inner lateral face and a postero-lateral costa on 
the outer lateral face. Group C is an asymmetrical 
multi-costate element (= slightly asymmetrical form 
group of Sweet and Bergstrom 1962) with a twisted 
cusp and two costae on each side. Group D is a tri- 
costate element, similar to group B but less laterally 
compressed with costa on the outer lateral face situated 
more towards the middle. Group E is a scandodiform 
element represented by the form species Scandodus 
unistriatus Sweet and Bergstrom 1962 (here assigned 
to the Pb position). 

Zhang (1998b) illustrated one of the multi- 
costate specimens as the P element, and two 
morphologically different scandodiform specimens as 
the M element. One of the latter specimens (Zhang 
1998b, pi. 15, fig. 19) is comparable with the form 
species S. unistriatus Sweet and Bergstrom 1962, and 
is regarded here as representing the Pb element of P. 
varicostatus (Fig. 80, P). The other specimen 
illustrated as the M element (Zhang 1998b, pi. 15, fig. 
14), which was recovered from the same sample with 
other illustrated specimens of P. varicostatus, has a 
multi-costate inner lateral face with three costae 
bordering two grooves and a few, shorter and weaker 
secondary costae near the base. It is designated here 
as occupying the Pa position of the species. 

Similar specimens (arcuatiform) referrable to 
the Pa element of P. varicostatus were also reported 
from allochthonous limestone clasts within the Shinnel 
Formation of Scotland (Armstrong 1997, pi. 3, figs 3, 
4). Morphologically it resembles the Pa element of 
Protopanderodus cf. calceatus Bagnoli and Stouge 
1996, recovered from the allochthonous limestones in 
the Oakdale Formation of central New South Wales 
(Zhen and Percival in press, pi. 17, figs A, C). 
However, this latter element has one larger, open 
groove on the inner lateral face, while the Pa element 
of P. varicostatus from South China (Zhang 1998b, 
pi. 15, fig. 14) and from the Wahringa area (Fig. 8N) 
has two equally developed, narrower grooves with a 
sharp costa between them. Protopanderodus liripipus 
Kennedy et al. 1979 is also multi-costate, but with a 
more posteriorly extended base (Fig. 8Y, Z). 

Three morphotypes of multicostate (S) 
elements with two costae on each side are recognised 
from sample CI 675 and possibly should be assigned 
to the Sd, Sb and Sc? positions, as no tri-costate 
elements have been recovered. The Sd element is 



Proc. Linn. Soc. N.S.W., 125, 2004 



157 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 



symmetrical with a reclined cusp and a short base. The 
Sb element is weakly asymmetrical with suberect cusp 
and a longer base. The Sc? element is asymmetrical 
and laterally strongly compressed with a suberect cusp 
and a short base. 

Genus PYGODUS Lamont and Lindstrom 1957 

Type species 

Pygodus anserinus Lamont and Lindstrom 1957. 

.Pygodus protoanserinus Zhang 1998a 
Fig. 9B-J 
Synonymy 
Pygodus anserinus Lamont and Lindstrom 1957, p. 

68, only fig. Id. 
Pygodus serrus (Hadding); Bergstrom 1971, p. 149, 

pi. 2, figs 22, 23; An 1981, pi. 4, figs 1-3; 

An 1987, pi. 24, fig. 25, pi. 26, figs 3, 6, 

13, pi. 29, figs 2, 3; Nicoll 1980, fig. 3H-L; 

Ding et al. in Wang, 1993, p. 198, pi. 30, 

figs 10, 13, 15-18, 20-22, 24, pi. 35, 24, 26. 
Pygodus protoanserinus Zhang 1998a, p. 96, Fig. 

2D, pi. 3, figs 9-18 (cum syn.); Zhang 

1998b, p. 86, 87, pi. 16, figs 6-8 (cum 

syn.). 
Pygodus serra (Hadding); Percival et al. 1999, fig. 

8.18; Pickett and Percival 2001, fig. 4C. 

Material 

Four Pa (stelliscaphate), five Pb (pastinate), and one 
Sb (tertiopedate) elements. 

Discussion 

Five species were assigned to Pygodus in the recent 
study of the genus by Zhang (1998a). They have short 
stratigraphic ranges and hence are very useful 
biostratigraphic index fossils. Sweet and Bergstrom 
(1962) and Bergstrom (1971) initially suggested that 
the apparatus of Pygodus anserinus, the type species 
of the genus, included elements represented by the 
form species Pygodus anserinus Lamont and 
Lindstrom 1957, and Haddingodus serrus (Hadding). 
Bergstrom (1971) also raised the possibility that the 
Pygodus apparatus might include elements represented 
by the form species Tetraprioniodus lindstroemi Sweet 
and Bergstrom 1962 and Roundya pyramidalis Sweet 
and Bergstrom 1962. This quadrimembrate Pygodus 
apparatus composition has been widely accepted 
(Ldfgren 1978, Clark et al. 1981, Sweet 1988). 
Subsequently, Armstrong (1997) has implied a 
septimembrate apparatus for Pygodus, but with only 
confirmed elements occupying the Pa, Pb, Pc, M and 
Sc positions. By including two pygodiform elements 
in the apparatus, Armstrong (1997) suggested that the 



P. anserinus apparatus consisted of the stelliscaphate 
Pa, pastiniscaphate Pb, bipennate Pc (= pastinate Pb 
of other authors' usage - see Zhang 1998a, b), 
tertiopedate M (termed an S element by other authors 
- see Zhang 1998a, b, and herein), and the ramiform 
Sc element. More recently Zhang (1998a, 1998b) 
proposed a quinquimembrate apparatus for Pygodus, 
including stelliscaphate Pa, pastinate Pb, and three 
ramiform S elements (alate, tertiopedate and 
quadriramate). 

Distinctions between P. serra and P. 
protoanserinus were discussed in detail by Zhang 
(1998a). Pygodus protoanserinus ranges from the E. 
robustus Subzone to the E. lindstroemi Subzone of the 
upper serra Zone in Baltoscandia, Scotland, North 
America, China, and Australia. The stelliscaphate Pa 
element from the lower part of the Wahringa Limestone 
Member is identical with the type material of P. 
protoanserinus, being characterised by having the 
middle denticle row situated more towards the outer 
denticle row on the upper surface. Specimens 
illustrated by Nicoll (1980) as P. serrus from the 
Pittman Formation at Black Mountain, Canberra, ACT, 
are here reassigned to P. protoanserinus on this same 
basis. A single specimen from the lower part of the 
Wahringa Limestone Member has the middle row of 
the denticles positioned centrally, and is therefore 
referred to P. serra (Fig. 9K, L), being more 
comparable with the middle form of the Pa element of 
that species as defined by Zhang (1998a). 

Genus STIPTOGNATHUS Ethington, Lehnert, and 
Repetski 2000 

Type species 

Reutterodus borealis Repetski 1982. 

Stiptognathus sp. A 
Fig. 90, P 
Synonymy 

Stiptognathus sp. Zhen and Percival in press, fig. 
21L-0. 

Material 

Two specimens from sample C1463 from an 
allochthonous limestone within the Fairbridge 
Volcanics, stratigraphically below the Wahringa 
Limestone Member. 

Discussion 

The symmetrical Sa and geniculate M elements 
recovered from sample CI 463 are identical with those 
from the allochthonous limestones of the Oakdale 
Formation (Zhen and Percival in press). Denticles on 
these specimens are small, closely spaced and blunt. 



158 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 




Figure 9. A, Pseudooneotodus mitratus (Moskalenko 1973); upper view, MMMC2701, C1474. B-J, Pygodus 
protoanserinus Zhang 1998a; B, Pa element, upper view, MMMC2702, C1450; C, Pa element, upper 
view, MMMC2703, C1652; D, upper view, and E, enlargement showing surface structure, Pa element, 
MMMC2704, C1652; F, inner lateral view, G, outer lateral view, H, anterior view, Pb element, 
MMMC2705, C1652; I, outer lateral view, J, showing surface structure, Sb element, MMMC2706, C1652. 
K, L, Pygodus serra (Hadding 1913); K, upper view, L, basal view, Pa element, MMMC2707, C1652. M, 
N, Stiptognathus sp. B; M, antero-lateral view, N, posterior view, Sa element, MMMC2708, C1675. O, P, 
Stiptognathus sp. A; O, M element, MMMC2709, C1463, posterior view; P, Sa element MMMC2710, 
C1463, antero-lateral view. Unless otherwise indicated scale bars are 100 urn. 



Proc. Linn. Soc. N.S.W., 125, 2004 



159 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 



Stiptognathus sp. B 
Fig. 9M, N 
Material 

Two specimens (Sa) from sample C1675. 

Discussion 

The cusp of this symmetrical element is triangular in 
cross section with a gently curved, wide anterior face, 
posterior costa, and an antero-lateral costa on each side. 
Three sharp costae extend basally into three blade-like 
processes, which bear small, upward-pointed denticles 
along the edges. They are easily distinguishable from 
the blunt denticles of Stiptognathus sp. A. 



Acknowledgments 

This study was supported by a Science Fellowship 
provided by the Sydney Grammar School to Zhen. Initial 
field collecting was undertaken with the support of the 
Australian Research Council during 1996 to 1999 (grant 
A39600788 to B.D. Webby). Gary Dargan from the 
Geological Survey of N.S.W. assisted in acid leaching, 
separation and other laboratory work. A grant to Y.Y. Zhen 
from the Betty Mayne Scientific Research Fund of the 
Linnean Society of New South Wales defrayed costs of some 
of the SEM work. The scanning electron microscope 
illustrations were prepared in the Electron Microscope Unit 
of the Division of Life and Environmental Sciences, 
Macquarie University and in the Electron Microscope Unit 
of the Australian Museum. I.G. Percival publishes with the 
permission of the Director General, New South Wales 
Department of Mineral Resources. 

REFERENCES 

Albanesi, G.L., Hiinicken, M.A. and Barnes, C.R. (1998). 
'Bioestratigrafia, biofacies y taxbnomia de 
conodontes de las secuencias Ordovicicas del 
cerro porterillo, Precordillera central de San 
Juan, R. Argentina' (ed. M.A. Hunicken). Actas 
de la Academia Nacional de Ciencias 12, 1-253. 

Armstrong, H.A. (1997). Conodonts from the Ordovician 
Shinnel Formation, southern Uplands, Scotland. 
Palaeontology 40, 763-797. 

An T.X. (1981). Recent progress in Cambrian and 

Ordovician conodont biostratigraphy of China. 
Geological Society of America Special Paper 
187, 209-226. 

An T.X. (1987). 'Early Paleozoic conodonts from South 
China'. 238 p. (Peking University Publishing 
House: Beijing) (in Chinese with English 
abstract). 

An, T.X., Zhang, F., Xiang, W.D., Zhang, Y.Q., Xu, 

W.H., Zhang, H.J., Jiang, D.B., Yang, C.S., Lin, 
L.D., Cui, Z.T. and Yang, X.C. (1983). 'The 
conodonts in North China and adjacent regions'. 



223 pp. (Science Press: Beijing) (in Chinese 
with English abstract). 

An, T.X. and Zheng, S.C. (1990). 'The conodonts of the 
marginal areas around the Ordos Basin, north 
China'. 199 pp. (Science Press: Beijing) (in 
Chinese with English abstract). 

Bagnoli, G. and Stouge, S. (1996). Lower Ordovician 

(Billingenian - Kunda) conodont zonation and 
provinces based on sections from Horns Udde, 
north Oland, Sweden. Bollettino della Societa 
Paleontologica Italiana 35, 109-163. 

Barnes, C.R. (1977). Ordovician conodonts from the Ship 
Point and Bad Cache Rapids Formations, 
Melville Peninsula, southeastern District of 
Franklin. Geological Survey of Canada, Bulletin 
269, 99-119. 

Bauer, J.A. (1987). Conodonts and conodont 

biostratigraphy of the McLish and Tulip Creek 
formations (Middle Ordovician) of south-central 
Oklahoma. Oklahoma Geological Survey, 
Bulletin 141, 1-55. 

Bauer, J.A. (1990). Stratigraphy and conodont 

biostratigraphy of the upper Simpson Group, 
Arbuckle Mountains, Oklahoma. In 'Early to 
Middle Paleozoic Conodont Biostratigraphy of 
the Arbuckle Mountains, Southern Oklahoma' 
(ed. S.M. Ritter). Oklahoma Geological Survey 
Guidebook 27, 39-46. 

Bauer, J.A. (1994). Conodonts from the Bromide 

Formation (Middle Ordovician), south-central 
Oklahoma. Journal of Paleontology 68, 358- 
376. 

Bergstrom, S.M. (1971). Conodont biostratigraphy of die 
Middle and Upper Ordovician of Europe and 
Eastern North America. Geological Society of 
America, Memoir 127, 83-157. 

Bergstrom, S.M. (1978). Middle and Upper Ordovician 
conodont and graptolite biostratigraphy of the 
Marathon, Texas graptolite zone reference 
standard. Palaeontology 21, 723-307. 

Bergstrom, S.M. (1990). Biostratigraphic and 

biogeographic significance of Middle and Upper 
Ordovician conodonts in the Girvan succession, 
south-west Scotland. Courier Forschungsinstitut 
Senckenberg 118, 1-43. 

Bergstrom, S.M., Carnes, J.B., Ethington, R.L., Votaw, 
R.B. and Wigley, P.B. (1974). 
Appalachignathus, a new multielement 
conodont genus from the Middle Ordovician of 
North America. Journal of Paleontology 48, 
227-235. 

Bergstrom, S.M. and Sweet, W.C. (1966). Conodonts 
from the Lexington Limestone (Middle 
Ordovician) of Kentucky and its lateral 
equivalents in Ohio and Indiana. Bulletin of 
American Paleontology 50 (229), 271-441. 

Branson, E.B. and Mehl, M.G. (1933). Conodont studies. 
University of Missouri Studies 8, 1-349. 

Clark, D.L., Sweet, W.C, Bergstrom, S.M., Klapper, G, 
Austin, R.L., Rhodes, F.H.T., Muller, K.J., 
Ziegler, W., Lindstrom, M., Miller, J.F. and 



160 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



Harris, A.G. (1981). Conodonta. In 'Treatise on 
Invertebrate Paleontology, part W, Miscellanea, 
supplement 2'. (ed. R.A. Robison). 202 pp. (The 
Geological Society of America, Boulder, and the 
University of Kansas, Lawrence). 

Dzik, J. (1976). Remarks on the evolution of Ordovician 
conodonts. Acta Palaeontologica Polonica 21, 
395-455. 

Dzik, J. (1978). Conodont biostratigraphy and 

paleogeographical relations of the Ordovician 
Mojcza Limestone (Holy Cross Mts, Poland). 
Acta Palaeontologica Polonica 23, 51-72. 

Dzik, J. (1991). Evolution of oral apparatuses in the 
conodont chordates. Acta Palaeontologica 
Polonica 36, 265-323. 

Dzik, J. (1994). Conodonts of the Mojcza Limestone. In 
'Ordovician carbonate platform ecosystem of 
the Holy Cross Mountains' (eds J. Dzik, E. 
Olempska, and A. Pisera). Palaeontologia 
Polonica 53, 43-128. 

Ethington, R.L. (1959). Conodonts of the Ordovician 

Galena Formation. Journal of Paleontology 33, 
257-292. 

Ethington, R.L. and Schumacher, D. (1969). Conodonts of 
the Copenhagen Formation (Middle Ordovician) 
in central Nevada. Journal of Paleontology 43, 
440-484. 

Ethington, R.L. and Clark, D.L. (1982). Lower and 

Middle Ordovician conodonts from the Ibex 
area, western Millard County, Utah. Brigham 
Young University Geology Studies 28 (2), 1-160. 

Ethington, R.L., Lehnert, O. & Repetski, J.E. (2000). 

Stiptognathus new genus (Conodonta: Ibexian, 
Lower Ordovician), and the apparatus of 
Stiptognathus borealis (Repetski, 1982). 
Journal of Paleontology 74, 92-100. 

Fahraeus, L.E., 1966. Lower Viruan (Middle Ordovician) 
conodonts from the Gullhogen Quarry, Southern 
Central Sweden. Sveriges Geologiska 
Undersbkning Ser. C 610, 1-40. 

Fahraeus, L.E. and Hunter, D.R. (1985a). Simple-cone 
conodont taxa from the Cobbs Arm Limestone 
(Middle Ordovician), New World Island, 
Newfoundland. Canadian Journal of Earth 
Sciences 22, 1171-1182. 

Fahraeus, L.E. and Hunter, D.R. (1985b). The curvature- 
transition series: integral part of some simple- 
cone conodont apparatuses (Panderodontacea, 
Distacodontacea, Conodontata). Acta 
Palaeontologica Polonica 30, 177-189. 

Glen, R.A., Walshe, J.L., Barron, L.M. and Watkins, J.J. 
(1998). Ordovician convergent-margin 
volcanism and tectonism in the Lachlan sector 
of east Gondwana. Geology 26, 751-754. 

Glenister, A.T. (1957). The conodonts of the Ordovician 
Maquoketa Formation in Iowa. Journal of 
Paleontology 31, 715-735. 

Hadding, A.R. (1913). Undre dicellograptusskiffern i 
Skane jamte nagra dagra darmed ekvivalenta 
bildningar. Lunds Universitets Arsskrift, Ny 
Foljd, Afdelning 2, 9(15), 1-90. 



Johnston, D.I. and Barnes, C.R. (2000). Early and Middle 
Ordovician (Arenig) conodonts from St. Pauls 
Inlet and Martin Point, Cow Head Group, 
western Newfoundland, Canada. 2. Systematic 
paleontology. Geologica et Palaeontologica 34, 
11-87. 

Kennedy, D.J., Barnes C.R. and Uyeno T.T. (1979). A 
Middle Ordovician conodont faunule from the 
Tetagouche Group, Camel Back Mountain, New 
Brunswick. Canadian Journal of Earth Sciences 
16, 540-551. 

Lamont, A. and Lindstrom, M. (1957). Arenigian and 
Llandeilian cherts identified in the Southern 
Uplands of Scotland by means of conodonts, 
etc. Transactions of the Edinburgh Geological 
Society 17, 60-70. 

Lehnert, O. (1995). Ordovizische Conodonten aus der 

Prakordillere Westargentiniens: Hire Bedeutung 
fur Stratigraphie und Palaogeographie. Erlanger 
Geologische Abhandlungen 125, 1-193. 

Lehnert, O., Bergstrom, S.M., Keller, M. and Bordonaro, 
O. (1999). Ordovician (Darriwilian-Caradocian) 
conodonts from the San Rafael Region, west- 
central Argentina: biostratigraphic, 
paleoecologic, and paleogeographic 
implications. In 'Studies on Conodonts - 
Proceedings of the Seventh European Conodont 
Symposium, Bologna-Modena, 1998' (ed. E. 
Serpagli). Bollettino della Societd 
Paleontologica Italiana 37, 199-214. 

Lee, H.Y. (1975). Conodonten aus dem unteren und 
mittleren Ordovizium von Nordkorea. 
Palaeontographica Abteilung A 150, 161-186. 

Leslie, S.A. (1997). Apparatus architecture of Belodina 
(Conodonta): Interpretations based on fused 
clusters of Belodina compressa (Branson and 
Mehl, 1933) from the Middle Ordovician 
(Turinian) Plattin Limestone of Missouri and 
Iowa. Journal of Paleontology 71, 921-926. 

Lin, B.Y., Qiu, H.R. and Xu, C.C. (1984). New 

observations of Ordovician strata in Shetai 
District of Urad Front Bannor, Nei Mongol 
(Inner Mongolia). Geological Review 30, 95- 
105. 

Lindstrom, M. (1955a). Conodonts from the lowermost 
Ordovician strata of south-central Sweden. 
Geologiska i Stockholm Foerhandlinger 76, 
517-604. 

Lindstrom, M. (1955b). The conodonts described by A.R. 
Hadding, 1913. Journal of Paleontology 29, 
105-111. 

Lindstrom, M. (1971). Lower Ordovician conodonts of 
Europe. In 'Symposium on conodont 
biostratigraphy' (eds W.C. Sweet and S.M. 
Bergstrom). Geological Society of America, 
Memoir 111, 21-61. 

Lofgren, A. (1978). Arenigian and Llanvirnian conodonts 
from Jamtland, northern Sweden. Fossils and 
Strata 13, 1-129. 

McCracken, A.D. (1987). Description and correlation of 
Late Ordovician conodonts from the D. ornatus 



Proc. Linn. Soc. N.S.W., 125, 2004 



161 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 



and P. pacificus graptolite zones, Road River 
Group, northern Yukon Territory. Canadian 
Journal of Earth Sciences 1A, 1450-1464. 

McCracken, A.D. (1989). Protopanderodus 

(Conodontata) from the Ordovician Road River 
Group, Northern Yukon Territory, and the 
evolution of the genus. Geological Survey of 
Canada Bulletin 388, 1-39. 

McCracken, A.D. (1991). Middle Ordovician conodonts 
from the Cordilleran Road River Group, 
northern Yukon Territory, Canada. In 
'Ordovician to Triassic conodont paleontology 
of the Canadian Cordillera' (eds M.J. Orchard 
and A.D. McCracken). Geological Survey of 
Canada, Bulletin 417, 41-63. 

McCracken, A.D. (2000). Middle and Late Ordovician 
conodonts from the Foxe Lowland of southern 
Baffin Island, Nunavut. In 'Geology and 
paleontology of the southeast Arctic Platform 
and southern Baffin Island' (eds A.D. 
McCracken and T.E. Bolton). Geological Survey 
of Canada, Bulletin 557, 159-216. 

McCracken, A.D. and Nowlan, G.S. (1989). Conodont 

paleontology and biostratigraphy of Ordovician 
carbonates and petroliferous carbonates on 
Southampton, Baffin, and Akpatok islands in 
the eastern Canadian Arctic. Canadian Journal 
of Earth Sciences 26, 1880-1903. 

Meakin, N.S. and Morgan, E.J. (compilers) (1999). 

'Dubbo 1:250 000 Geological Sheet SI/55-4, 
2nd edition. Explanatory Notes'. (Geological 
Survey of New South Wales: Sydney). 

Moskalenko, T.A. (1972). Ordovician conodonts of the 
Siberian Platform and their bearing on 
multielement taxonomy. Geologica et 
Palaeontologica 1, 47-56. 

Moskalenko, T.A. (1973). Conodonts of the Middle and 
Upper Ordovician on the Siberian Platform. 
Akademiy Nauk SSSR, Sibirskoe Otdelenie, 
Trudy Instituta Geologii i Geofiziki 137, 1-143. 
(in Russian). 

Moskalenko, T.A. (1983). Conodonts and biostratigraphy 
in the Ordovician of the Siberian Platform. 
Fossils and Strata 15, 87-94. 

Nicoll, R. S. (1980). Middle Ordovician conodonts from 
the Pittman Formation, Canberra, ACT. BMR 
Journal of Australian Geology and Geophysics 
5, 150-153. 

Nowlan, G.S. (1981). Some Ordovician conodont 

faunules from the Miramichi Anticlinorium, 
New Brunswick. Geological Survey of Canada 
Bulletin 345, 1-34. 

Nowlan, G.S. and Barnes, C.R. (1981). Late Ordovician 
conodonts from the Vaureal Formation, 
Anticosti Island, Quebec; Part 1. Geological 
Survey of Canada, Bulletin 329, 1-49. 

Nowlan, G.S., McCracken, A.D. and Chatterton, B.D.E. 
(1988). Conodonts from Ordovician-Silurian 
boundary strata, Whittaker Formation, 
Mackenzie Mountains, Northwest Territories. 



Geological Survey of Canada, Bulletin 373, 1- 
99. 

Palmieri, V. (1978). Late Ordovician conodonts from the 
Fork Lagoons Beds, Emerald area, central 
Queensland. Geological Survey of Queensland 
Publication 369, Palaeontological Paper 43, 1- 
55. 

Pander, C. H. (1856). 'Monographic der fossilen Fische 

des Silurischen Systems der Russisch-Baltischen 
Gouvernements' . 91 pp. (Akademie der 
Wissenschaften, St. Petersburg). 

Percival, I.G., Morgan, E.J and Scott, M.M. (1999). 

Ordovician stratigraphy of the northern Molong 
Volcanic Belt: new facts and figures. Geological 
Survey of New South Wales, Quarterly Notes 
108, 8-27. 

Percival, I.G., Webby, B.D. and Pickett, J.W. (2001). 
Ordovician (Bendigonian, Darriwilian to 
Gisbornian) faunas from the northern Molong 
Volcanic Belt of central New South Wales. 
Alcheringa 25, 211-250. 

Philip, G.M. (1966). The occurrence and 

palaeogeographic significance of Ordovician 
strata in northern New South Wales. Australian 
Journal of Science 29, 112-113. 

Pickett, J.W. and Percival, I.G. (2001). Ordovician faunas 
and biostratigraphy in the Gunningbland area, 
central New South Wales, Australia. Alcheringa 
25, 9-52. 

Pohler, S.M.L. (1994). Conodont biofacies of Lower to 
lower Middle Ordovician megaconglomerates, 
Cow Head Group, Western Newfoundland. 
Geological Survey of Canada, Bulletin 459, 1- 
71. 

Pyle, L.J. and Barnes, C.R. (2002). 'Taxonomy, evolution, 
and biostratigraphy of conodonts from the 
Kechika Formation, Skoki Formation, and Road 
River Group (Upper Cambrian to Lower 
Silurian), Northeastern British Columbia'. 227 
pp. (NRC Research Press, Ottawa). 

Rasmussen, J.A. (2001). Conodont biostratigraphy and 
taxonomy of the Ordovician shelf margin 
deposits in the Scandinavian Caledonides. 
Fossils and Strata 48, 1-180. 

Repetski, J.E. (1982). Conodonts from El Paso Group 
(Lower Ordovician) of westernmost Texas and 
southern New Mexico. New Mexico Bureau of 
Mines and Mineral Resources, Memoir 40, 1- 
121. 

Sansom, I.J., Armstrong, H.A. and Smith, M.P. (1995). 
The apparatus architecture of Panderodus and 
its implications for coniform conodont 
classification. Palaeontology 37, 781-799. 

Schopf, T.J. (1966). Conodonts of the Trenton Group 
(Ordovician) in New York, southern Ontario, 
and Quebec. New York State Museum and 
Science Service Bulletin 405, 1-105. 

Stauffer, C.R. (1935). The conodont fauna of the Decorah 
Shale (Ordovician). Journal of Paleontology 9, 
596-620. 



162 



Proc. Linn. Soc. N.S.W., 125, 2004 



Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 



Stouge, S. (1984). Conodonts of the Middle Ordovician 

Table Head Formation, western Newfoundland. 

Fossils and Strata 16, 1-145. 
Sweet, W.C. (1979). Late Ordovician conodonts and 

biostratigraphy of the western Midcontinent 

Province. Brig ham Young University Geology 

Studies 26, 45-86. 
Sweet, W.C. (1988). "The Conodonta: Morphology, 

Taxonomy, Paleoecology, and Evolutionary 

History of a Long-Extinct Animal Phylum'. 212 

pp. (Clarendon Press, Oxford). 
Sweet, W.C. and Bergstrom, S.M. (1962). Conodonts 

from the Pratt Ferry Formation (Middle 

Ordovician) of Alabama. Journal of 

Paleontology 36, 1214-1252. 
Tipnis, R.S., Chatterton, B.D.E. and Ludvigsen, R. 

(1978). Ordovician conodont biostratigraphy of 

the southern district of Mackenzie, Canada. In 

'Western and Arctic Canadian biostratigraphy' 

(eds C.R. Stelck and B.D.E. Chatterton). 

Geological Association of Canada, Special 

Paper 18, 39-91. 
Trotter, J.A. and Webby, B.D. (1995). Upper Ordovician 

conodonts from the Malongulli Formation, 

Cliefden Caves area, central New South Wales. 

AGSO Journal of Australian Geology and 

Geophysics 15 (4), 475-499. 
Wang, C.Y. (ed.) (1993). 'Conodonts of the Lower 

Yangtze Valley - an index to biostratigraphy and 

organic metamorphic maturity'. 326 pp. 

(Science Press, Beijing), (in Chinese with 

English Summary). 
Watson, S.T. (1988). Ordovician conodonts from the 

Canning Basin (W. Australia). 

Palaeontographica, Abteilung A 203 (4-6), 91- 

147. 
Webers, G.F., 1966. The Middle and Upper Ordovician 

conodont faunas of Minnesota. Minnesota 

Geological Survey, Special Publication 4, 1- 

123. 
Zhang, J.H. (1998a). The Ordovician conodont genus 

Pygodus. In 'Proceedings of the sixth European 

conodont Symposium (ECOS VI)' (ed. H. 

Szaniawski). Palaeontologia Polonica 58, 87- 

105. 
Zhang, J.H. (1998b). Conodonts from the Guniutan 

Formation (Llanvirnian) in Hubei and Hunan 

Provinces, south-central China. Stockholm 

Contributions in Geology 46, 1-161. 
Zhao, Z.X., Zhang, G.Z. and Xiao, J.N. (2000). 'Paleozoic 

stratigraphy and conodonts in Xinjiang'. 340 pp. 

(Petroleum Industry Press, Beijing) (in Chinese 

with English Abstract). 
Zhen, Y.Y. and Webby, B.D. (1995). Upper Ordovician 

conodonts from the Cliefden Caves Limestone 

Group, central New South Wales, Australia. 

Courier Forschungsinstitut Senckenberg 182, 

265-305. 



Zhen, Y.Y., Webby, B.D. and Barnes, C.R. (1999). Upper 
Ordovician conodonts from the Bowan Park 
succession, central New South Wales, Australia. 
Geobios 32, 73-104. 

Zhen, Y.Y., Nicoll, R.S., Percival, I.G., Hamedi, MA. 
and Stewart, I. (2001). Ordovician 
rhipidognathid conodonts from Australia and 
Iran. Journal of Paleontology 75, 186-207. 

Zhen, Y.Y., Percival, I.G. and Farrell, J.R. (2003a). Late 
Ordovician allochthonous limestones in Late 
Silurian Barnby Hills Shale, central western 
New South Wales. Proceedings of the Linnean 
Society of New South Wales 124, 29-51. 

Zhen, Y.Y., Percival, I.G. and Webby, B.D. (2003b). 

Early Ordovician conodonts from western New 
South Wales, Australia. Records of the 
Australian Museum 55, 169-220. 

Zhen, Y.Y., Percival, I.G. and Webby, B.D. (in press). 

Early Ordovician (Bendigonian) conodonts from 
central New South Wales. Courier 
Forschungsinstitut Senckenberg 245. 

Zhen, Y.Y. and Percival, I.G. (in press). Middle 

Ordovician (Darriwilian) conodonts from 
allochthonous limestones in the Oakdale 
Formation of central New South Wales. 
Alcheringa 28. 

Ziegler, W. (ed.) (1981). 'Catalogue of Conodonts, Vol. 
4'. 445 pp. (Schweizerbart'sche Verlags- 
buchhandlung, Stuttgart). 



Proc. Linn. Soc. N.S.W., 125, 2004 



163 



ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 

APPENDIX 
LOCALITY DATA 

All grid references are AMG66 co-ordinates and relate to the Cumnock 8632-S 1:50,000 topographic sheet 
(first ed., 1978). 

Allochthonous limestones stratigraphically below Wahringa Limestone Member 
C1463: GR 679150 mE 6371900 mN 
C1486: GR 678750 mE 6371800 mN 
C1487: GR 678720 mE 6371800 mN 
C1488: GR 678700 mE 6371800 mN 

Wahringa Limestone Member, type section 

C1450, C1652 (basal beds): GR 678650 mE 6372000 mN 

C1456, C1664, C1667-C1668, C1672 (middle beds): GR 678700 mE 6372050 mN 

C1458, C1464, C1673-1683, C1687 (upper beds): GR 678700 mE 6372100 mN 

Wahringa Limestone Member, northeast extremity of outcrop 
C1707, C1709-1713: centred on GR 679330 mE 6372620 mN 

Wahringa Limestone Member, southwest extremity of outcrop (middle or upper beds) 
C1429: GR 677570 mE 6370800 mN 

Allochthonous limestones stratigraphically above Wahringa Limestone Member 
C1693, C1694: GR 678000 mE 6371310 mN 
C1695, C1696: GR 678100 mE 6372000 mN 
C1697, C1698: GR 678050 mE 6372000 mN 
CI 699: GR 678025 mE 6372000 mN 
C1700: GR 678010 mE 6372000 mN 
C1471: GR 678500 mE 6372200 mN 
C1472: GR 679250 mE 6373550 mN 
C1474: GR 678100 mE 6372600 mN 
C1483: GR 678500 mE 6372800 mN 



164 Proc. Linn. Soc. N.S.W., 125, 2004 



Wenlock (Early Silurian) Brachiopods from the Orange District 

of New South Wales 

A.J. Wright 1 and D.L. Strusz 2 

'School of Earth and Environmental Sciences, University of Wollongong, Wollongong NSW 2522, 
tony_wright@uow.edu.au; department of Earth and Marine Sciences, Australian National University, 

Canberra ACT 0200; dstrusz@ems.anu.edu.au 



Wright, A.J. and Strusz, D.L. (2003). Wenlock (Early Silurian) brachiopods from the Orange District of 
New South Wales. Proceedings of the Linnean Society of New South Wales 125, 165-172. 



Two late Wenlock (Early Silurian) brachiopod species from the Ulah Formation near Orange, New 
South Wales, are closely associated with graptolite faunas. Visbyella cumnockensis occurs in the testis 
Biozone on Wallace Creek in the Four Mile Creek area, and Strophochonetes melbournensis is recorded 
from the ludensis Biozone on Spring Creek. Poorly preserved but similar Visbyella! and Strophochonetes! 
From the Prfdoli Wallace Shale at Cheesemans Creek are also illustrated. These occurrences provide significant 
new stratigraphic and distributional data for the species. 

Manuscript received 19 March 2003, accepted for publication 22 October 2003. 

KEYWORDS: Brachiopods, New South Wales, P0fdoll, Silurian, Strophochonetes melbournensis, Ulah 
Formation, Visbyella cumnockensis, Wallace Shale, Wenlock. 



INTRODUCTION 

The Silurian strata of the area west and 
southwest of Orange, NSW, in the valleys of Spring 
Creek and Four Mile Creek (Fig. 1), have yielded a 
diversity of fossils, but very few shelly fossils have 
ever been described, apart from corals described by 
authors including Etheridge and McLean (full 
references to these works can be found in Pickett 1982). 
The most abundant and important fossils in the region 
are graptolites, which have been known for more than 
50 years and were reported by Packham and Stevens 
(1955) and Jenkins (1978, 1986). 

Jenkins recorded (but did not describe) 
brachiopod faunas from limestones in the Four Mile 
Creek area, but few brachiopods have been reported 
from clastic strata common in the area. Rickards and 
Wright (1997) described two brachiopod species from 
late Wenlock strata (ludensis Biozone) in Cobblers 
Creek (Fig. 1), and in the section at 'Mirrabooka Park' 
brachiopods were noted in Wenlock strata during field 
work by L. Muir, R.B. Rickards, G.H. Packham and 
A.J. Wright. A diverse and abundant shelly fauna 
occurring with the late Wenlock graptolite 



Testograptus testis on the Cadia gold mine access road, 
several kilometres to the east of Four Mile Creek, was 
illustrated by Rickards et al. (2001). 

The two species described here are recorded 
for the first time from the region near Orange. One, 
Visbyella cumnockensis Walmsley et al., 1968, was 
originally described from near Cumnock, 55 km 
northwest of Orange, where it occurs with T. testis 
(Walmsley et al. 1968:315). Visbyella has been 
reported also, but not illustrated, by Pickett (1982) and 
Pogson and Watkins (1998). The other species, 
Strophochonetes melbournensis (Chapman 1903), was 
previously known only from Wenlock and Ludlow 
strata in the Melbourne Trough, Victoria. Pickett's 
report was based on the record of Visbyella cf. 
cumnockensis by Sherwin (1971). Sherwin's locality 
is younger, and contains a sparse and poorly preserved 
fauna including also a chonetoide similar to 
Strophochonetes? savagei Strusz, 2000 from 
Cumnock. These taxa are illustrated but not described. 
Documented brachiopod occurrences in the Orange 
region are still insufficient, however, to permit any 
notion of a regional brachiopod zonation. 



EARLY SILURIAN BRACfflOPODS FROM ORANGE NSW 




Figure 1. Map of the area southwest to west of Orange, central 
New South Wales, showing the geographic context of the two 
localities, LM3 on Wallace Creek east of Cargo and W940 near 
'Mirrabooka Park' east-southeast of Cudal. Inset: the location 
of Orange within Australia. 



Wenlock to Pridoli; the age of the strata 
at this locality is late Wenlock. 

W940. 

The somewhat more abundant 
specimens of Strophochonetes 
melbournensis were collected from 
dark siltstones of the Ulah Formation 
on the southern side of Spring Creek 
at 'Mirrabooka Park' , directly opposite 
One Tree Hill. There are also 
occasional poorly preserved 
brachiopods, including pentamerides, 
in beds at about the same level on One 
Tree Hill itself. The shells at W940 
occur with a graptolite fauna that 
includes Monograptus ludensis (R.B. 
Rickards, pers. comm.). Only 
disarticulated valves are known at this 
locality; small phosphatic brachiopods 
are quite common, and there are rare 
specimens of other brachiopods 
including strophomenides and 
atrypides. Most specimens of 
Strophochonetes melbournensis at this 
locality retain shelly material and the 
spines on the pedicle valve hinge line 
are often preserved. The environment 
was most probably a low-energy one. 



LOCALITIES 

LM3. 

Visbyella cumnockensis was collected from 
Wallace Creek in the Four Mile Creek area, in grey- 
brown siltstones assigned by Jenkins (1978) to the 
Wenlock-Ludlow Ulah Formation. These beds have 
also yielded the graptolites Cyrtograptus and a new 
species of Monograptus (L. Muir, pers. comm.), and 
overlie beds containing T. testis. The brachiopod 
specimens are moulds of a single pedicle and a single 
brachial valve on the same bedding surface, which 
could represent the disarticulated valves of a single 
shell. No other macrofossils have been found at this 
locality. In contrast, the type material of V. 
cumnockensis is entirely of specimens in the 'butterfly' 
position, with the shell opened so that the conjoined 
valves lie on the bedding surface. The age assigned to 
the Ulah Formation by Chapman et al. (2003) is late 



MO/I/27. 

A few poorly preserved orthide 
and chonetoide specimens have been 
collected from this outcrop of fine thin- 
bedded siltstone low in the Wallace 
Shale, about 600 m east of 'Mirrabooka' homestead. 
The fauna also includes occasional trilobites. The lo- 
cality lies within the Monograptus transgrediens 
Biozone. 

SYSTEMATIC PALAEONTOLOGY 

Suprageneric taxonomy follows that in Kaesler 
(2000); references to authorship of suprageneric taxa 
are therefore not repeated here. Specific diagnoses have 
been rephrased to accord with currently accepted 
terminology (Kaesler 1997). Details of localities are 
given in the descriptive section below. 

Abbreviations. 

Ls - shell length 
Ld - dorsal valve length. 
Ws - shell width 
Wh - hinge line width 



166 



Proc. Linn. Soc. N.S.W., 125, 2004 



A.J. WRIGHT AND D.J. STRUSZ 



Figure 2. a-g, Visbyella cumnockensis Walmsley et 
al., 1968. a-c, ventral valve counterparts; a, latex 
cast of exterior, AM F124331. b-c, internal mould 
and latex cast, AM F124332. d-g, dorsal valve 
counterparts; d, latex cast of exterior, AM F124333. 
e-g, internal mould and latex cast (in ventral and 
postero-ventral views), AM F124334. h-k, cf. 
Visbyella cumnockensis, Pridoli, Wallace Shale, h, 
latex cast of ventral valve, MM F37431. i, latex cast 
of shell in 'butterfly' position, MM F21132. j, 
external mould of dorsal valve plus internal mould 
of ventral valve, MM F21125. k, latex cast of 
incomplete interior of shell in 'butterfly' position, 
MM F37428. Scale bar 2 mm. 



AM - Australian Museum 

MM - Mining Museum Collection, 

Geological Survey of NSW 
CPC - Commonwealth Palaeontological 

Collection 
NMV - Museum of Victoria 
SU - Sydney University (Geology 

Department) 



Suborder DALMANELLIDINA Moore 1952 
Superfamily DALMANELLOIDEA Schuchert 1913 
Family DALMANELLJDAE Schuchert 1913 
Subfamily RESSERELLINAE Walmsley and 
Boucot 1971 

Genus VISBYELLA Walmsley, Boucot, Harper and 
Savage 1968 

Type species 

Orthis visbyensis Lindstrom 1861, by original 
designation; late Llandovery, Gotland. 

Diagnosis 

Subcircular, small valves with apical deltidium 
and hypercline dorsal interarea; ventral interior with 
recessive dental plates and cordate muscle scar; dorsal 
interior with trilobed, dorsally-facing cardinal process 
and median septum (Harper p. 797 in Kaesler 2000). 



Visbyella cumnockensis Walmsley, 

Boucot, Harper and Savage 1968 

Fig. 2 (a-g) 

Synonymy 

1968 Visbyella cumnockensis sp. nov. 
Walmsley et al., pp. 313-315, pi. 61 figs 6-12. 




Type material 

Holotype AM F67781; paratypes AM F67782- 
67788 (formerly SU P19511, 19512-19518; all 
renumbered when collections were transferred from 
the University of Sydney to the Australian Museum). 

New material 

External and internal moulds of a ventral valve 
(AM F124331, F124332) and a dorsal valve (AM 



Proc. Linn. Soc. N.S.W., 125, 2004 



167 



EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW 



F124333, F124334) from one bedding plane at locality 
LM3 (Grid reference 782 988, Cudal 8631 II and III 
50 000 topographic sheet, Wallace Creek, Four Mile 
Creek area south of Orange, N.S.W.); Ulah Formation, 
Testograptus testis Biozone; late Wenlock (Early 
Silurian). 

Diagnosis 

Relatively small, weakly sulcate, coarsely 
multicostellate Visbyella with semicircular outline. 
Dorsal median ridge broad and low posteriorly, 
becoming narrower and higher to form an anterior 
median septum (after Walmsley et al. 1968) 

Description 

Shell small, almost plano-convex. Ventral valve 
broadly naviculate, with low suberect beak; dorsal 
valve weakly convex with shallow but distinct sulcus. 
Outline suboval, moderately transverse, with straight 
hinge, obtuse slightly rounded cardinal angles; greatest 
width at about 0.4Ls. Ventral interarea strongly 
apsacline, almost flat, apical angle about 120°; 
delthyrium open, apical angle about 70°, rimmed by 
narrow crescentic deltidium. Dorsal interarea low, 
concave, catacline, apical angle about 150°; 
notothyrium filled by cardinal process, apical angle 
about 80°. Ribs rather angular, stronger medially than 
laterally, increasing by bifurcation on the ventral valve, 
intercalation on the dorsal valve; about 30 counted at 
ventral valve margin. 

Ventral interior with prominent subtriangular 
muscle field, impressed posteriorly but slightly raised 
anteriorly, length l/3Ls and width l/4Ws. Diductor 
scars elongate oval, divergent, depressed a little below 
slightly shorter flat adductor field. Raised anterior 
margin to adductor field distinctly denticulate, extends 
forward to about 3/4Ls as low ridge. Vascula media 
flank this ridge as broad, shallow furrows extending 
from the diductor scars. Muscle field flanked by stout 
dental plates, divergent forward at about 100° and 
slightly divergent ventrally, not extending beyond 
muscle field. Teeth strong, wide, triangular, with 
distinct crural fossettes on antero-median faces. Valve 
floor faintly radially furrowed, marginally strongly 
crenulated. 

Dorsal interior with prominent oval muscle 
field extending to 2/3Ld, width l/3Ws, defined by 
strong ridges arising just in front of brachiophores and 
increasingly raised anteriorly, which converge to abut 
on median septum. Diductor scars impressed, elongate 
oval, subequal, posterior scars subparallel, anterior 
scars convergent forward; scars separated by tapering 
ridge from which rises the stout median septum. 
Septum highest a little in front of muscle field, and 
extends to valve margin. Cardinal process large, 



directed posterodorsally, continuous with well 
developed notothyrial platform; no shaft. 
Brachiophores stout, blade-like, divergent ventrally, 
supported by low, thick plates. Sockets oval, diverging 
from valve axis at about 75°, deeply excavated into 
thick triangular socket pads. Valve floor radially 
grooved, marginally strongly crenulated. 



Dimensions 








AM F124332 


AM F124334 


valve 


ventral 


dorsal 


Ls, Ld 


est 2.85 


2.59 


Ws 


3.90* 


3.73 


Wh 


3.60* 


3.32 


Ls/Ws 


est. 0.73 


0.69 


Wh/Ws 


est. 0.92 


0.89 



* values obtained by doubling exposed half-width, 
assuming a symmetrical shell. 

Remarks 

The Wallace Creek occurrence of this species 
is almost exactly the same age as the original 
occurrence at Cumnock, and our admittedly limited 
new material corresponds closely in all specific 
characters to the type material. The specimens are 
slightly larger than shells of the type series (the 
maximum length and width of any specimens of the 
type series are 2.1 mm and 3.1 mm respectively), but 
the ratio Ls:Ws is close to the 2:3 cited for the type 
material; while the marginal crenulations in the ventral 
valve are less extensive. The internal moulds of the 
disarticulated valves are somewhat better than the 
types, and features of the hinge line can be seen more 
clearly. 

The species was also tentatively recorded by 
Sherwin (1971, p. 223) from the Pridolf Wallace Shale 
at locality MO/I/27 in the Cheesemans Creek area north 
of Quarry Creek; his report was the basis for 
subsequent reports by Pickett (1982, pp. 154-155) and 
Pogson and Watkins (1998, p. 131). This occurrence 
is in significantly younger strata than the two other 
occurrences noted herein. Sherwin' s report was based 
on several specimens from one locality; we were 
recently guided to this locality by Dr Sherwin, and 
collected a further seven specimens of the 'orthid' 
species, which is very rare at the locality (also collected 
were a few poor specimens of a chonetide, identified 
as Strophochonetesl cf. savagei Strusz, 2000, and 
illustrated in Fig. 4 for comparison with 
Strophochonetes melbournensis). 

Unfortunately the only internal mould of a 
dorsal valve of the Wallace Shale orthide (Fig. 2k) is 
incomplete, and appears to lack a median septum, 
although its presence anteriorly cannot be completely 
ruled out. It was initially thought that the absence of a 



168 



Proc. Linn. Soc. N.S.W., 125, 2004 



A.J. WRIGHT AND D.J. STRUSZ 



septum would rule out the presence of Visbyella. 
However, one specimen (AM F 125552) of Visbyella 
cumnockensis on one of the type slabs is close in size 
to the Wallace Shale material and, unlike all the other 
type specimens, lacks a median septum, so this is not 
an infallible character of this species. Other 
morphological features of the Wallace Shale material 
are not well preserved; there appear to be more than 
30 costellae, and the internals of both valves, in so far 
as they are preserved, are similar to those of the 
Wallace Creek material (compare Figs 2h-i with Fig. 
2a, and Fig. 2j with Fig. 2b). 

Hence no conclusive argument can be presented 
to refute the presence of Visbyella at this locality, 
unlikely as it might seem. This opinion is slightly 
supported by the presence of a similar orthide 
(probably Resserella), but definitely lacking a median 
septum, in the late Ludlow Cardinal View Shale (Bauer 
1994) at Bungonia, NSW. Unfortunately, our 
experience gives us no reason to expect more definitive 
material at this very unproductive Wallace Shale 
locality. 



Suborder CHONETIDINA Muir-Wood 1955 
Superfamily CHONETOIDEA Bronn 1862 
Family STROPHOCHONETIDAE Muir-Wood 1962 
Subfamily STROPHOCHONETINAE Muir-Wood 

1962 
Genus STROPHOCHONETES Muir-Wood 1962 

Type species 

Chonetes cingulatus Lindstrom 1861, by 
original designation; Wenlock, Gotland. 



1945 Chonetes (Chonetes) melbournensis 

Chapman; Gill, pp. 132-133. 

1953 Chonetes infantilis n. sp.; Opik; p. 15, 

pi. m, figs 19-22. 

2000 Strophochonetes melbournensis 

(Chapman, 1903); Strusz, pp. 249-251, figs 

2-3. 

Type material 

Lectotype NMV P1419, paralectotypes NMV 
P615-6, 619, 623, 625-7, 630-633, 637-43 designated 
by Strusz (2000); Melbourne Formation, Melbourne 
and South Yarra, Victoria; Ludlow (Late Silurian). 
Type material of Chonetes infantilis Opik, 1953: 
holotype CPC 661, paratypes CPC 662-663, Illaenus 
Band, Wapentake Formation, Heathcote, Victoria; late 
Wenlock (Early Silurian). 

New material 

AM F124306 - 124330, locality W940 (grid 
reference 743 123, Cudal 8631-11 and III 50 000 
topographic sheet; south bank of Spring Creek, 
'Mirrabooka Park', southwest of Orange, central 
N.S.W.); Ulah Formation, with Monograptus ludensis; 
Late Wenlock (Early Silurian). 

Diagnosis 

Small, weakly concavo-convex, subquadrate 
Strophochonetes with up to 5 pairs of gently intraverse- 
cyrtomorph hinge spines, and finely multicostellate 
ornament with median rib on ventral valve usually 
strongly enlarged. Valve floors heavily papillose, 
ventral muscle field distinct, anderidia short and 
diverging at 60-80° (after Strusz 2000). 



Diagnosis 

Shell small, piano- to moderately concavo- 
convex; well developed median enlarged costa; long, 
symmetrically arranged high-angled spines varying 
from intraverse cyrtomorph proximally to orfhomorph 
vertical distally; cardinal process strongly bilobed 
internally, anteriorly bounded by cardinal process pit; 
no median septum; anderidia long, narrow, anteriorly 
divergent at about 60° and isolated on valve floor; inner 
socket ridges short, thin, as two rounded ridges almost 
parallel to hinge (after Racheboeuf p. 369 in Kaesler 
2000). 



Strophochonetes melbournensis (Chapman 1903) 
Fig. 3 

Synonymy 

1903 Chonetes melbournensis sp. nov.; 
Chapman, pp. 74-76, pi. XI, fig. 2 only. 



Description 

Shell small, plano-convex, ventral valve of very 
low convexity. Outline subquadrate, lateral margins 
gently sigmoid, with shallow re-entrants in front of 
small triangular ears; hinge width usually slightly less 
than greatest width (mean Wh/Ws 0.93). Ventral 
protegulum posteromedially furrowed, variably raised 
above remaining shell surface; distinct protegular lobe, 
weaker lateral lobes on dorsal valve. Maximum 
observed width 9.8 mm, length 6.5 mm, most 
specimens being much smaller; mean Ls/Ws 0.75, ratio 
decreasing with increasing shell size. Interareas mostly 
obscure; ventral interarea apparently low, apsacline, 
flat, delthyrium wide, beak very low; pseudodeltidium 
not seen; dorsal interarea linear, attitude unclear. 
Myophore small, projecting posteroventrally, bifid, 
each lobe less strongly bifid, flanked by small but 
distinct cardinal crests. Chilidium obscure, might be 
present as very narrow ridge wrapped around base of 
myophore. Hinge spines fine, relatively long, upright 



Proc. Linn. Soc. N.S.W., 125, 2004 



169 



EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW 




Figure 3. Strophochonetes melbournensis (Chapman, 1903). a-g, ventral valves; some hinge spines are 
visible in b-e, only spine bases in f-g. a, juvenile valve AM F124320. b, juvenile with particularly prominent 
protegulum, AM F124317. c, juvenile AM F124322. d, AM F124324. e-f, internal mould and latex cast, 
AM F124312. g, AM F124326. h-j, dorsal valves; h-i, incomplete external mould and latex cast showing 
well developed protegular and lateral nodes, AM F124318. j, latex cast of incomplete interior, AM F124307. 
Scale bar 3 mm. 



or nearly so (initial angle with hinge line about 60- 
80°), straight (particularly in small specimens) to gently 
cyrtomorph intraverse, symmetrically placed; up to 4 
each side of beak (AM F 124324). Ornament of fine, 
rounded radial ribs, 29-34 counted in 5 mm at 5 mm 
radius, separated by narrower furrows; increase is by 
bifurcation only. Median rib on ventral valve 
prominent, arises within protegulum; remaining ribs 
arise at or in front of margins of concentrically 
wrinkled protegular regions. 

Ventral interior with low, narrow median 
septum, reaching forward to about 0.2Ls; septum 
posteriorly raised and slightly widened. Teeth small, 
widely divergent. Muscle field generally obscure other 
than for weak or absent endospines; in one specimen 
(AM F124312) the field is weakly impressed, with 



small, elongate subtriangular, slightly divergent 
adductor scars further impressed posteriorly. 
Remainder of valve floor densely covered by fine 
endospines radially arranged beneath ribs, weakest 
towards cardinal margin and ears. 

Dorsal interior still not well known. Cardinal 
process small, internally bifid, fused to short but strong 
inner socket ridges which are curved parallel to hinge 
margin. Short, shallow furrow in front of cardinal 
process, but no median ridge developed. Anderidia 
visible in only one specimen (AM F124307); they are 
short (0.2Ld), fine, low, diverging at about 60°. Muscle 
field obscure. Distal two-thirds of valve floor with 
numerous small radially arrayed endospines, as in 
ventral valve. 



170 



Proc. Linn. Soc. N.S.W., 125, 2004 



AJ. WRIGHT AND D J. STRUSZ 



Dimensions 
















valve 


Ls,Ld 


Ws 


Wh 


Ls/Ws 


Wh/ 1 


AM F 124307 


dorsal 


4.9 


- 


8.7* 


- 


- 


F124312 


ventral 


est. 4.8 


5.6* 


5.4* 


est. 0.86 


0.96 


F124318 


dorsal 


5.5 


- 


- 


- 


- 


F124322 


ventral 


3.6 


4.7 


4.4 


0.77 


0.94 


F124324 


ventral 


5.3 


7.2 


7.2 


0.74 


1.00 


F124326 


ventral 


5.6 


8.4 


7.2 


0.67 


0.86 



* values obtained by doubling exposed half- width, assuming a symmetrical shell. 



Discussion 

Although preservation is not particularly good, 
the Wenlock specimens from Spring Creek conform 
in all important aspects (very low ventral convexity, 
rib increase only by bifurcation, and less prominent 
protegular and lateral lobes on the dorsal valve) with 
S. melbournensis rather than S. kemezysi Strusz, 2000. 
Some of the minor differences could be related to the 
small size of most of the specimens (several are clearly 
juvenile, none approaches the maximum size recorded 
for the Victorian material). Some could be of age 
significance, but without better and more abundant 
material from older levels in Victoria this remains 
unclear. Thus no ventral valves show the anterior 
sulcus seen in some Victorian Late Silurian shells, and 
no more than 4 spines have been seen to either side of 
the ventral beak. The NSW specimens tend also to be 
more elongate (Ls/Ws very variable, mean 0.76; for 
Victorian specimens the mean is 0.61). Internally, the 
ventral muscle field is less obvious, and there are no 
coarser endospines near the hinge. In this last respect, 
and in a greater tendency for spines on small specimens 
to be straight, the Late Wenlock Spring Creek 
specimens are more like the few poor specimens from 
the Early Wenlock of Heathcote than the Ludlow 
material from Melbourne. Dorsal interiors, while still 
few and inadequate, do add some information, 




3 mm 



Figure 4. Strophochonetes? cf. savagei Strusz, 2000. 
Latex cast of ventral valve, MM F21133. Scale bar 
3 mm. 



particularly the form of the cardinal process and its 
flanking cardinal crests. The presence internally of a 
weak posteromedian dorsal furrow instead of a low 
ridge places these specimens closer to typical 
Strophochonetes than are the type specimens. 

Three similar chonetoide specimens (MM 
F21 133, 37435, 37436) are available from the Wallace 
Shale locality - the best of them is figured (Fig. 4). All 
are small and weakly convex. In the absence of internal 
data, particularly of the dorsal valve, generic identity 
must remain uncertain. The long more or less upright 
hinge spines, low ventral valve convexity, fine ribbing 
and accentuated median rib all indicate 
Strophochonetes, however, and of the Australian taxa 
described by Strusz (2000) the closest is undoubtedly 
S? savagei from the Early Lochkovian of Manildra, 
northwest of Orange. S. melbournensis and S. kemezysi 
Strusz, 2000, while superficially similar, are both larger 
and more coarsely ribbed; the latter has very prominent 
protegulae. In only one respect these specimens appear 
unlike typical Strophochonetes, and that is in the 
alternating pattern of hinge spine insertion described 
for instance by Strusz (2000, p. 259) for the strongly 
convex and fairly coarsely ribbed Australian species 
of Johnsonetes Racheboeuf, 1987 (all of which lack 
spine 1'). However it is not clear that spine 1' is 
undeveloped in the Wallace Shale specimens. 
Moreover, the Manildra species show considerable 
variation in spine form, and some asymmetry cannot 
be ruled out. 



ACKNOWLEDGMENTS 

We gratefully acknowledge access graciously made 
available by Ian Street to 'Mirrabooka Park' and Ken 
Williams to 'Ashburnia', and thank Dr L. Sherwin for 
drawing our attention to the report of Visbyella cf. 
cumnockensis from the Wallace Shale and subsequently 
guiding us to the locality. Prof. Barrie Rickards and Dr Lucy 
Muir kindly allowed us to cite identifications of the 
graptolites. Robert Jones readily made the type material of 
Visbyella cumnockensis available for study. Strusz wishes 
to thank Dr Patrick DeDeckker for providing facilities at the 



Proc. Linn. Soc. N.S.W., 125, 2004 



171 



EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW 



Australian National University; Wright's research has been 
supported by the University of Wollongong and the Linnean 
Society of NSW. 



REFERENCES 

Bauer, J. A. (1994). Siluro-Devonian Bungonia Group, 
Southern Highlands, N.S.W. Helictite 32(2), 25-34. 

Chapman, A.J., Rickards, R.B., Wright, A.J., and Packham, 
G.H. (2003). Dendroid and tuboid graptolites from 
the Llandovery (Silurian) of the Four Mile Creek 
. area, New South Wales. Records of the Australian 
Museum. 

Chapman, F. (1903). New or little-known Victorian fossils 
in the National Museum. Part II - some Silurian 
Molluscoidea. Proceedings of the Royal Society of 
Victoria 16, 60-82. 

Gill, E.D. (1945). Chonetidae from the Palaeozoic rocks of 
Victoria and their stratigraphical significance. 
Proceedings of the Royal Society of Victoria 57, 125- 
150. 

lenkins, C.J. (1978). Llandovery and Wenlock stratigraphy 
of the Panuara area, central New South Wales. 
Proceedings of the Linnean Society of New South 
Wales 102, 109-130. 

Jenkins, C.J. (1986). The Silurian of mainland Australia: a 
field guide. (IUGS Silurian Subcommission and 
University of Sydney: Sydney). 

Kaesler, R.L. (ed.) (1997). Treatise on Invertebrate 
Paleontology, PartH, Brachiopoda, revised, volume 
1: Introduction. (Geological Society of America and 
University of Kansas Press: Lawrence, Kansas). 

Kaesler, R.L. (ed.) (2000). Treatise on Invertebrate 
Paleontology, Part H, Brachiopoda, revised, 
volumes 2-3: Linguliformea, Craniiformea, and 
Rhynchonelliformea (part). (Geological Society of 
America and University of Kansas Press: Lawrence, 
Kansas). 

Lindstrom, G. (1861). Bidrag till kannedomen om Gotlands 
brachiopoder. Ofversigt af kungliga Vetenskaps- 
Akademiens Forhandlingar, Stockholm for 1 860, 17, 
337-382. 



Muir-Wood, H. (1962). On the morphology and 
classification of the brachiopod suborder 
Chonetoidea. (British Museum of Natural History: 
London). 

Opik, A.A. (1953). Lower Silurian fossils from the "Illaenus 
Band", Heathcote, Victoria. Geological Survey of 
Victoria, Memoir 19, 1-42. 

Packham, G.H., and Stevens, N.C. (1955). The Palaeozoic 
stratigraphy of Spring and Quarry Creeks, west of 
Orange, N.S.W. Journal and Proceedings of the 
Royal Society of New South Wales 88, 55-60. 

Pickett, J.W. (ed.) 1982. The Silurian System in New South 
Wales. Geological Survey of New South Wales, 
Bulletin 29,1- 264. 

Pogson, D.J., and Watkins, J.J. (compilers) 1998. 'Bathurst 
1: 250 000 Geological Sheet S 1/55-8: Explanatory 
Notes'. (Geological Survey of New South Wales: 
Sydney). 

Racheboeuf, P.R. (1987). Upper Lower and Lower Middle 
Devonian chonetacean brachiopods from Bathurst, 
Devon and Ellesmere Islands, Canadian Arctic 
Archipelago. Geological Survey of Canada, 
Bulletin 375, 1-29. 

Rickards, R.B., Percival, I.G., Simpson, A.J. and Wright, 
A.J. (2001). Silurian biostratigraphy of the Cadia 
area, near Orange, New South Wales. Proceedings 
of the Linnean Society of New South Wales, 123, 
173-191. 

Rickards, R.B., and Wright, A.J. (1997). Graptolite zonation 
of the late Wenlock, with a new graptolite- 
brachiopod fauna from New South Wales. Records 
of the Australian Museum 49, 229-248. 

Sherwin, L. (1971). Stratigraphy of the Cheesemans Creek 
district, New South Wales. Records of the Geological 
Survey of New South Wales 13, 199-237. 

Strusz, D.L. (2000). Revision of the Silurian and Early 
Devonian chonetoidean brachiopods of southeastern 
Australia. Records of the Australian Museum 52, 
245-287. 

Walmsley, V.G, Boucot, A.J., Harper, C.W. and Savage, 
N.M. (1968). Visbyella - a new genus of resserellid 
brachiopod. Palaeontology 11, 306-316. 



172 



Proc. Linn. Soc. N.S.W., 125, 2004 



Early Silurian Graptolites from Cadia, New South Wales 

R.B. Rickards 1 AND A. J. Wright 2 

'Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, England 
and 2 School of Earth and Environmental Sciences, University of Wollongong, Wollongong NSW 2522. 

Rickards, R.B. and Wright, A.J. (2004). Early Silurian graptolites from Cadia, New South Wales. 
Proceedings of the Linnean Society of New South Wales 125, 173-175. 

A low-diversity graptolite fauna is reported from the Ulah Formation at Cadia, central western New South 
Wales. The assemblage includes Testograptus testis, Monoclimacis flumendosae, fragments of Monograptus 
flemingii, possible Cyrtograptus and unidentifiable retiolitid meshworks, and is correlated with the lungreni- 
testis Biozone, of late Wenlock (Early Silurian) age. 

Manuscript received 3 May 2003, accepted for publication 23 July 2003. 

KEY WORDS: Cadia, graptolites, Lower Silurian, Wenlock. 



INTRODUCTION 

Three Silurian faunas were documented by 
Rickards et al. (2000) from the vicinity of Cadia open 
cut, south of Orange, New South Wales. One of these 
faunas, of late Wenlock-early Ludlow aspect, consisted 
of shelly fossils and graptolites collected by Dr Ian 
Percival from a slumped mudstone at a locality on the 
access road to the Cadia open cut. This fauna was 
discussed and illustrated by Rickards et al. (2000), who 
figured but could not determine the poor graptolite 
material to genus or species because of the poor 
preservation of the fragmentary material. The locality 
(grid reference 687240E, 6295047 N, Canowindra 
8360-N 1:50 000 topographic sheet) is on the eastern 
face of the access road to the Cadia open cut, about 1 
km from the entrance gates; a map of the region 
showing the location of this and other fossil localities 
was provided by Rickards et al. (2000, Fig. 1). The 
fossiliferous strata are considered to correlate with the 
Ulah Formation, at Four Mile Creek west of Cadia (see 
Rickards et al. 2000, Fig. 1), in which the Testograptus 
testis fauna occurs. 



NOTES ON THE GRAPTOLITE FAUNA 



Monoclimacis flumendosae (Gortani), fragments of 
Monograptus flemingii (Salter), fragmentary stipes 
possibly belonging to Cyrtograptus, and fragmentary 
retiolitid meshworks which cannot be assigned, even 
approximately, to a genus. 

In discussing this as 'the Cadia graptolite 
fauna' we are mindful of the presence of other 
graptolites in Silurian strata in the vicinity of the Cadia 
mine. Full documentation of any such graptolite faunas 
as that documented here is important as graptolite 
localities in the vicinity of Cadia mine (such as the 
Pridoli 'borrow pit' locality, W910 of Rickards et al. 
2000) are very much less common than at Four Mile 
Creek, and are under threat. A brief review of 
graptolites previously reported from Cadia by 
Offenberg (1963) was given by Rickards et al. (2000). 

We have not provided here any systematic 
descriptions of the fauna, but limited comments on 
the morphological detail are included in the 
explanatory text for Figure 1. The Cadia specimens 
have undergone soft sediment deformation, with a 
considerable amount of twisting and breakage, in 
contrast to the Rodds Creek black shale specimens 
(Rickards et al. 2000) which were undeformed other 
than by diagenetic flattening. 



Since the publication of Rickards et al. 
(2000), we have made a further but small graptolite 
collection from the Cadia mine shelly fossil locality 
which permits fuller identification of the low-diversity 
fauna and determination of its age. The Cadia graptolite 
fauna consists of Testograptus testis (Barrande), 



AGE OF THE CADIA GRAPTOLITE FAUNA 

The dominant species is Testograptus testis 
(Barrande), which normally indicates the late Wenlock 
(Early Silurian) lundgreni-testis Biozone. Testograptus 
testis has been recorded, very rarely, from the ludensis 



SILURIAN GRAPTOLITES FROM NEW SOUTH WALES 




Figure 1. (A) Monoclimacis flumendosae (Gortani), AM Fl 14926, distal thecae, undeformed, low relief. 
(B-E) Testograptus testis (Barrande). (B) proximal end, AM F114928, showing some soft sediment 
deformation distally; (C) AM F114925, a proximal end with spines visible on thl; (D) AM F114930, 
spines on several thecae; (E) AM Fl 14929, distal thecae with a growing end visible. (F) Monograptus 
flemingii (Salter), AM Fl 14927, subscalariform view of mesial thecae. 

AH figures xlO, scale bar 1mm; heavy bar indicates deformation stretching direction, possibly not tectonic. 
All specimens from locality W 937, grid reference 687240E, 6295047 N, Canowindra 8360-N 1:50 000 
topographic sheet. Unfigured specimens are AMF 114931-940. 



Biozone (Rickards et al. 1995) but, as the Cadia 
specimens are abundant and occur with Monoclimacis 
flumendosae (Gortani), a pre-ludensis Biozone is 
indicated for this fauna. 

The Cadia fauna is probably slightly younger 
than the Rodds Creek fauna (Rickards et al. 2000). 
Although this latter assemblage included some 
lundgreni-testis Biozone indicators, the presence of 
Cyrtograptus ex gr. rigidus Tullberg indicated a 
probable middle rather than late Wenlock for the Rodds 
Creek fauna. The Cadia fauna is thus significantly older 
than the Pridolf fauna from the 'borrow pit' locality 
(W910) 2 km to the southeast (Rickards et al. 2000). 



Correlation with the Four Mile Creek 
sequence is probably with testis-beaiing beds of the 
Ulah Formation in Wallace Creek; in Spring and 
Quarry Creeks, the tons-bearing beds of the same 
formation are largely green and black mudstones 
(Packham, Rickards and Wright, unpublished data). 



SHELLY FAUNAS 

The disarticulated and fragmental shelly 
fauna in this slump unit is unusually abundant and 
diverse for the region, in contrast with clastic units of 



174 



Proc. Linn. Soc. N.S.W., 125, 2004 



R.B. RICKARDS AND A.J. WRIGHT 



this age in the Four Mile Creek area and the Spring- 
Quarry Creek areas which are singularly poor in shelly 
fossils. The faunas at Cadia have undergone soft- 
sediment deformation and are clearly transported. 
Described shelly faunas (other than corals) from the 
Four Mile Creek area and the Spring Creek areas are 
limited to two species of ludensis Biozone brachiopods 
described by Rickards and Wright (1997) from 
Cobblers Creek (see Fig. 1 of Rickards et al. 2000) 
and by Wright and Strusz (2004) from Spring Creek 
and Wallace Creek (see Fig. 1 of Rickards et al. 2000: 
ludensis Biozone and lundgreni-testis Biozone 
respectively). Other bracbiopod faunas from the region 
were listed by Jenkins (1978, 1986), but the only rich 
faunas cited by him are from Llandovery (Early 
Silurian) limestones. 



CONCLUSIONS 

Graptolites identified from the Cadia Mine 
access road locality are Testograptus testis, 
Monoclimacis flumendosae, fragments of 
Monograptus flemingii, ICyrtograptus and retiolitids. 
The fauna is late Wenlock (Early Silurian) and is 
probably best correlated with a level high in the 
lundgreni-testis Biozone. It appears to be slightly 
younger than the probably middle Wenlock Rodds 
Creek black shale fauna (Rickards et al. 2000), and is 
assumed to correlate with the testis fauna of the Ulah 
Formation in the Four Mile Creek area to the west of 
Cadia. 



REFERENCES 

Jenkins, C.J. (1978). Llandovery and Wenlock stratigraphy 
of the Panuara area, central New South Wales. 
Proceedings of the Linnean Society of New South 
Wales 102, 109-130. 

Jenkins, C.J. (1986). The Silurian of mainland Australia: a 
field guide. 82 p. IUGS Silurian Subcommission, 
Sydney. 

Offenberg, A.C. (1963). Geology of the Panuara-Cadia- 
Errowanbong area, south of Orange, New South 
Wales. BSc (Hons) thesis, University of Sydney 
(unpublished), 112 p. 

Rickards, R.B., Packham, G.H., Wright, A.J. and 

Williamson, PL. (1995). Wenlock and Ludlow 
graptolite faunas and biostratigraphy of the Quarry 
Creek district, New South Wales. Association of 
Australasian Palaeontologists, Memoir 17, 1-68. 

Rickards, R.B., Percival, I.G., Simpson, A.J. and Wright, 
A.J. (2000). Silurian biostratigraphy of the Cadia 
area, near Orange, New South Wales. Proceedings 
of the Linnean Society of New South Wales 123, 
173-191. 

Rickards, R.B. and Wright, A.J. (1997). Graptolite 

zonation in the late Wenlock (Early Silurian), with 
a new graptolite-brachiopod fauna from New 
South Wales. Records of the Australian Museum 
49, 229-248. 

Wright, A.J. and Strusz, DL. (2004). Wenlock (Early 

Silurian) brachiopods from the Orange district of 
New South Wales. Proceedings of the Linnean 
Society of New South Wales 125, 165-172. 



ACKNOWLEDGEMENTS 

We are grateful to Ian Tedder (Newcrest Mining 
Limited) for allowing access to, and guiding us to, this 
locality in November 2001. The universities of Cambridge 
and Wollongong have provided financial support for 
participation by RBR and AJW in this study, and funds were 
provided to AJW by the Linnean Society of New South 
Wales through the Betty Mayne Fund. 



Proc. Linn. Soc. N.S.W., 125, 2004 



175 



176 



Silicified Early Devonian Trilobites from Brogans Creek, New 

South Wales 

Gregory D. Edgecombe 1 and Anthony J. Wright 2 

'Australian Museum, 6 College Street, Sydney, NSW 2010 (greged@austmus.gov.au); 
2 School of Earth and Environmental Sciences, University of Wollongong, Wollongong, NSW 2522 

(tony_wright @ uow.edu.au) 

Edgecombe, G.D. and Wright, A.J. (2004). Silicified Early Devonian trilobites from Brogans Creek, New 
South Wales. Proceedings of the Linnean Society of New South Wales 12A, 177-188. 

Trilobites in an Emsian silicified fauna from the Carwell Creek Formation at Brogans Creek SE of Mudgee, 
NSW, include Acanthopyge (Jasperia) bifida, Dentaloscutellum hudsoni and Proetus nemus, all originally 
described from the Taemas area of NSW, together with Sthenarocalymene. Proetus nemus was known from 
limited material at Taemas, but is the most abundant species at Brogans Creek. Fuller description substantiates 
membership in Proetus {-Devonoproetus), rather than Ryckholtia, Longiproetus or Rhenocynproetus. Early 
ontogenetic stages of the trilobites are lacking at Brogans Creek, in contrast to Taemas. Conodonts co- 
occurring with the shelly fauna at Brogans Creek and at Taemas include Polygnathus nothoperbonus, which 
indicates the Polygnathus perbonus Conodont Zone (medial Emsian). 

Manuscript received 16 October 2003, accepted for publication 8 January 2004. 

Keywords: Trilobita, Devonian, New South Wales, Acanthopyge {Jasperia), Dentaloscutellum, Proetus, 
Sthenarocalymene. 



INTRODUCTION 

The presence of Devonian limestone at 
Brogans Creek (Fig. 1), located SE of Mudgee in the 
central tablelands of NSW, was first noted by Carne 
and Jones (1919) and later by Lishmund et al. (1986). 
Fossils from the limestone were discussed in detail by 
Colquhoun (1998) and Colquhoun and Meakin in 
Colquhoun et al. (in Meakin and Morgan 1999). 
Colquhoun (1995) illustrated the conodonts 
Pandorinellina e. exigua and Polygnathus 
nothoperbonus from Brogans Creek, the latter species 
considered (after Mawson 1987) to be characteristic 
of the medial Emsian {Polygnathus perbonus zone). 

Here we provide the first descriptions of any 
of the well-preserved and abundant fossils from the 
quarry at Brogans Creek. A silicified trilobite fauna is 
of low diversity, but it provides new data on some taxa 
described from the Taemas area by Chatterton (1971), 
in particular the proetid Proetus nemus. 

Stratigraphic assignment and age 

The Devonian strata at Brogans Creek were 
considered part of the Carwell Creek Formation by 
Colquhoun et al. (1999). The limestones that have 
yielded the trilobites and other fossils documented here 
have also yielded (Colquhoun 1995) the medial Emsian 



conodont Polygnathus nothoperbonus, so this 
limestone is significantly younger than most limestones 
occurring in the area between the Mudgee and Brogans 
Creek, with the principal exception of those reported 
by Pickett (1972) and Colquhoun (1998) from the 
Mount Knowles Limestone Member of the Carwell 
Creek Formation and by Pickett ( 1 97 8) from the Mount 
Frome Limestone, both located to the E of Mudgee. 

Little is known about the sequence of the 
Devonian strata in the vicinity of the Brogans Creek 
quarry, and recent land reclaimation operations have 
concealed formerly productive parts of the abandoned 
quarries. Colquhoun (1998) stated that the sequence 
grades upwards from the fossiliferous limestone 
through crinoidal sandstone into massive shale and 
volcarenite. The sequence of beds that yielded silicified 
fossils is about 10 m in thickness. Beds immediately 
overlying these strata have yielded the tetracorals 
Xystriphyllum mitchelli and Embolophyllum, both also 
described from the Receptaculites Limestone Member 
at Taemas and Wee Jasper by Pedder et al. (1970). 

The similarity of the macrofauna to that from 
the Receptaculites and Warroo limestone members of 
the Taemas Formation in the Burrinjuck Dam area of 
NSW (see Pedder et al. 1970) necessitates some 
consideration of the ages of these units. Conodont data 
summarised by Talent et al. (2000) for the Taemas 



EARLY DEVONIAN TRILOBITES FROM N.S.W. 





Sofala 

5 10 km 



Figure 1. Location of trilobite collection at Brogans Creek. Map of NSW indicates Taemas, where the 
same species have been described (Chatterton 1971). Shading in inset map (after Colquhoun 1995) shows 
distribution of Lower Devonian platform sediments. 



178 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE AND A.J. WRIGHT 



area of NSW indicate that the Receptaculites and 
Warroo limestones at Taemas, which overlie the Cavan 
Formation with its Polygnathus pireneae to P. 
dehiscens fauna, are probably early Emsian. Lindley 
(2002) recorded Polygnathus nothoperbonus from the 
Warroo Limestone Member, further confirming the 
assignment of this limestone to the medial Emsian 
Polygnathus perbonus Conodont Zone. However, 
Basden et al. (2002) concluded that the Warroo 
Limestone should be correlated with the Polygnathus 
inversus to P. serotinus Conodont Zones. On balance 
the co-occurrence of P. nothoperbonus in both areas 
of NSW seems to indicate unequivocally a medial 
Emsian age for the macrofaunas. This supports the 
conclusions of Garratt and Wright (1988:Fig. 3), who 
correlated their Malurostrophia-Taemostrophia- 
Howittia fauna (essentially the shelly fauna discussed 
here) with the Polygnathus gronbergi (=P. perbonus) 
Conodont Zone. 

Faunal characters and affinities 

The fossiliferous limestones have yielded 
very rich and well-preserved invertebrate faunas, 
dominated by brachiopods, tabulate corals and 
tetracorals, trilobites, gastropods, ostracodes, 
cephalopods, tentaculitids, crinoid debris and sponges; 
bivalves are subordinate at this locality. Most of the 
trilobites and brachiopods at Brogans Creek are 
conspecific with those described from Emsian 
limestones in the Lake Burrinjuck sequence at Taemas 
and 'Bloomfield' by Chatterton (1971, 1973). With 
respect to the trilobites, the faunal composition of the 
Brogans Creek assemblage is best matched in the lower 
half of the Receptaculites Limestone at Locality T of 
Chatterton (1971). The three species identified here, 
Proetus nemus, Dentaloscutellum hudsoni and 
Acanthopyge bifida, are represented in the lower 
Receptaculites Limestone at Locality T and at that 
locality as well as Brogans Creek they occur with 
Sthenarocalymene. Silicified residues from Brogans 
Creek yield the following for minimal number of 
individuals per species, based on the most abundant 
skeletal element: Proetus nemus (N=54), 
Dentaloscutellum hudsoni (N=16), Acanthopyge bifida 
(N=7), and Sthenarocalymene sp. (N=2). About 120 
kilograms of limestone have been etched to produce 
our fauna. 

In terms of diversity, the silicified assemblage 
consists additionally of more than 15 brachiopod 
species (Malurostrophia flabellicauda reverta 
Chatterton; Salopina kemezysi Chatterton and other 
dalmanellids; Schuchertella murphyi Chatterton; 
Coelospira dayi Chatterton; Howellella sp.; 
Ambothyris runnegari Chatterton; Howittia sp.; 



IBuchanathyris sp.; reticulariid indet.; Cydimia parva 
Chatterton; Parachonetes flemingi Chatterton; P. sp. 
cf. P. konincki Chatterton; rhynchonellids). Some 30 
gastropod species are under study by Dr A.G. Cook. 
Tetracoral species are dominated numerically by an 
abundant solitary Plasmophyllum, as well as other 
solitary corals (?acanthophyllids) and rare fragments 
of ICalceola. The sponge Amphipora is locally 
abundant, and presumably represents lagoonal phases 
of deposition or influx of lagoonal debris; several 
biofacies are evident. Colquhoun (1998) indicated that 
the Brogans Creek limestone was deposited in a well- 
oxygenated, normal salinity environment. The trilobite 
material is represented by disarticulated sclerites, but 
many brachiopods shells are articulated. Scolecodonts 
are at least as common as conodonts in residues; this 
is also a feature of limestones in the Capertee Valley 
(S of Brogans Creek) where the strata are highly 
deformed and preservation is poor. Despite the 
disarticulated nature of parts of the Brogans Creek 
shelly fauna, their excellent preservation indicates that 
postmortem transportation was minimal. 



SYSTEMATIC PALAEONTOLOGY 

Figured material is in the Palaeontology collection, 
Australian Museum, Sydney (prefix AMF). 

Order PROETIDA Fortey and Owens, 1975 

Family PROETIDAE Salter, 1864 

Subfamily PROETINAE Salter, 1864 

Genus PROETUS Steininger, 1831 

Type species 

Calymmene concinna Dalman, 1827; by original 
designation. 

Proetus nemus Chatterton, 1971 
Fig. 2a-p, Fig. 3a-t 

Proetus nemus Chatterton, 1971:65-67, PL 16, Figs 

18-32. 

Ryckholtial nemus (Chatterton). Lutke, 1990:21. 

Material 

39 cranidia, 103 librigenae, 3 hypostomes, 62 
thoracic segments, 50 pygidia. 

Diagnosis 

Proetus with relatively elongate, tapering glabella, its 
posterior two thirds with dense, mostly moderate sized 
tubercles, its anterior third granulate. Facial suture 
divergent between y and p\ Genal ridge strong along 



Proc. Linn. Soc. N.S.W., 125, 2004 



179 



EARLY DEVONIAN TRILOBITES FROM N.S.W. 




Figure 2. Proetus nemus Chatterton, 1971. Carwell Creek Formation (medial Emsian), Brogans Creek, 
NSW. Scale bars 1 mm. a-c, AMF 124700, cranidium, dorsal, anterior and lateral views; d-f, AMF 124701, 
cranidium, dorsal, anterior and lateral views; g, AMF 124702, cranidium, dorsal view; h, AMF 124703, 
cranidium, dorsal view; i, AMF 124704, cranidium, lateral view; j, AMF 124705, cranidium, anterior 
view; k, AMF 124706, cranidium, dorsal view; 1-m, AMF 124707, cranidium, dorsal and lateral views; n, 
AMF 124708, librigena, dorsal view; o, AMF 124709, librigena, dorsal view; p, AMF 125485, cranidium, 
dorsal view. 



180 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE AND A.J. WRIGHT 




Figure 3. Proetus nemus Chatterton, 1971. Carwell Creek Formation (medial Fmsian), Brogans Creek, 
NSW. Scale bars 1 mm. a, AMF 124710, librigena, internal view; b, AMF 124711, librigena, dorsal view; 
c, AMF 124712, librigena, dorsal view; d-e, AMF 124713, hypostome, ventral and lateral views; f, AMF 
124714, thoracic segment, dorsal view; g, AMF 124715, thoracic segment, dorsal view; h-j, AMF 124716, 
pygidium, posterior, lateral and dorsal views; k, AMF 124717, thoracic segment, dorsal view; 1, AMF 
124718, thoracic segment, anterior view; m, AMF 124719, thoracic segment, anterior view; n, AMF 124720, 
pygidium, dorsal view; o-p, AMF 124721, pygidium, lateral and dorsal views; q-r, AMF 124722, pygidium, 
posterior and dorsal views; s, AMF 124723, pygidium, ventral view; t, AMF 124724, pygidium, dorsal 
view. 



Proc. Linn. Soc. N.S.W., 125, 2004 



181 



EARLY DEVONIAN TRILOBITES FROM N.S.W. 



all but posteriormost part of librigenal field, distinct 
but less prominent on preocular fixigena; small caecal 
pits abundant on librigenal field; genal spine relatively 
long. Pygidium with seven axial rings and lunate 
terminal piece (7+1); anterior three or four pleural 
furrows well impressed, fifth and sixth faint. 

Description 

Cranidial length about equal to maximum width at co; 
width at 8 slightly more than 80% width at co; width at 
P 85-95% width at 8. Axial furrow narrow, moderately, 
evenly deep. Glabella widest basally, length (excluding 
L0) 1.1-1.2 times basal width, with moderate taper 
anteriorly, slightly constricted at S2, gently convex 
(sag., tr.); frontal lobe rounded; terminating at but not 
overhanging anterior border furrow. SI originating 
opposite midlength of palpebral lobe, shallow, directed 
posteromedially, distally birfucate, with posterior 
branch terminating well in front of SO; S2 parallel with 
SI, more weakly incised, originating just behind 
anterior edge of palpebral lobe; S3 obscure. Posterior 
two thirds of glabella with mostly moderate sized 
tubercles, some small tubercles, densely packed so as 
to nearly touch; anterior third of glabella granulate, 
non-tuberculate. SO transverse medially, narrow (sag., 
exsag.), deep, flexed forwards abaxially against lateral 
occipital lobes. L0 distinctly wider than basal part of 
glabella, length about 20% its width; lateral occipital 
lobes large, drop-shaped, isolated from remainder of 
L0 by deep furrows; L0, including lateral lobes, 
covered with tubercles as on posterior part of glabella, 
including moderately large median tubercle behind 
midlength. Preglabellar region 13-15% of cranidial 
length; in large specimens, composed of an inclined, 
medially flat posterior half and moderately convex 
(sag.) anterior half bearing 5-6 terrace lines in dorsal 
view; in small specimens, posterior half forms a wide 
(sag., tr.) depressed field with a broad (tr.), gently 
inflated transverse median swelling. Genal ridge well 
developed on preocular fixigena, anteromedially 
directed, terminating at juncture of preglabellar and 
anterior border furrows, stronger in small specimens. 
Postocular fixigena 25-35% width (tr.) and about 60% 
length (exsag.) of L0. Palpebral lobe arcuate, 35-45% 
length of glabella; palpebral furrow faint or indistinct. 
Anterior sections of facial suture diverging from each 
other at 45-62° between y and P, running subparallel 
against anterior border furrow, then strongly 
converging between P and a. Posterior sections of 
facial suture running subparallel or gently diverging 
between e and £, close to axial furrow, then sharply 
turned outwards to co. 

Librigenal field moderately wide, gently 
convex (tr.); genal ridge strong along all but 



posteriormost part of field, closer to eye socle than to 
lateral border furrow; most of field with abundant, 
small caecal pits, least distinct at posterolateral corner 
of field. Eye socle narrow, separated from visual 
surface and librigenal field by shallow furrows. 
Posterior border furrow narrow, deep; lateral border 
furrow wider, the two merging at genal angle, 
extending along a variable extent of the genal spine, 
usually along about half its length. Lateral border 70- 
80% as wide as narrowest part of librigenal field in 
dorsal view, strongly convex (tr.); terrace lines well 
defined along entire length and width of lateral border 
and along genal spine. Genal spine relatively long, its 
inner margin straight or faintly concave. Panderian 
notch large, semicircular. Connective suture with 
straight, diagonal course along most of its length, its 
extent relative to cranidium indicating that rostral plate 
is trapezoidal or triangular, fairly wide anteriorly (cf. 
P. concinnus: Owens 1973:Text-fig. IB). 

Hypostomal width across shoulders about 
65% sagittal length. Anterior margin weakly convex 
medially, flexed backward abaxially. Anterior lobe of 
middle body strongly inflated (tr.), anteromedial part 
raised but not forming discrete rhynchos; middle body 
gently convex (sag.) along most of length, fairly steeply 
turned up anteromedially; anterior lobe bearing many 
sinuous terrace lines. Middle furrow moderately deep, 
directed posterolaterally across abaxial third of middle 
body then abruptly effacing. Border furrow narrow, 
distinctly impressed around entire middle body, 
shallowest against anterior wing. Anterior border 
uniformly narrow (sag., exsag.); lateral border gently 
converging between anterior wing and shoulder; 
shoulder rounded; posterolateral margin straight 
between shoulder and pair of blunt spines at lateral 
edge of posterior border; posterior border narrow (sag., 
exsag.), about 10% length of hypostome, with gently 
convex posteromedian margin. 

Number of thoracic segments unknown. 
Axial furrow narrow, shallow. Axis strongly convex 
(tr.), 32-41% width of thorax. Articulating half ring 
varying from equal in width (sag.) to 1.6 times as wide 
as preannulus along length of thorax, 70-90% length 
of ring; preannular furrow transverse to gently concave 
medially, sharply impressed but much shallower than 
articulating furrow; ring covered with small, dense 
tubercles or coarse granules. Pleural furrow narrow, 
about as deep as articulating furrow, gently flexed 
forward at fulcrum, abruptly shallowing then effacing 
on inner part of articulating facet; anterior and posterior 
pleural bands equal in width (exsag.) proximal to 
fulcrum; pleurae moderately declined abaxial to 
fulcrum, at midwidth (tr.) of rib. Pleural tips with 
curved anterolateral margin, blunt rounded posterior 



182 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE AND A.J. WRIGHT 



projection. Panderian notch deep, U-shaped. 

Pygidium subsemicircular, length (excluding 
articulating half ring) 55-60% width. Axial width about 
35% pygidial width anteriorly; axial furrows narrow, 
uniformly impressed along most of length. Seven axial 
rings and short, lunate terminal piece (7+1); first one 
or two ring furrows lengthened medially as short 
preannulus; more posterior ring furrows shallower but 
with moderately deep incision across axis, posterior 
few gently convex backwards; axis raised strongly 
above pleurae, gently convex in sagittal profile, 
moderately arched (tr.); rings with dense small 
tubercles or coarse granules. Postaxial region about 
20% length of pygidium. Pleural furrows narrow 
(exsag.), anteriorly convex, anterior three or four well 
impressed, fifth and variably sixth faintly discernible; 
first pleural furrow terminates near pygidial lateral 
margin, others terminate at shallow posterior border 
furrow; interpleural furrows narrower and shallower 
than pleural furrows; pleural ribs with sculpture of 
dense, medium sized granules. Border widening back 
to its intersection with third pleural furrow, then 
maintaining even width, occupying most of postaxial 
region, weakly convex. Doublure extending in nearly 
as far as border furrow, bearing several terrace lines. 

Discussion 

The sample from Brogans Creek resembles that from 
Taemas in that the largest cranidia (Fig. 2a-c, h, i, p; 
Chatterton 1971:P1. 16, fig. 28) have the anterior end 
of the glabella abutting the inclined posterior part of 
the anterior border, whereas small specimens have a 
broad depression between the frontal lobe and the 
convex, terraced part of the anterior border (Fig. 2d-f, 
j, k; Chatterton 1971:P1. 16, fig. 25). The latter 
morphology, associated with a more pronounced 
fixigenal ridge (Fig. 2d, k versus 2a, h, p) is confined 
to small specimens. This difference in the preglabellar 
region is bridged by intermediate sized specimens, and 
is ascribed to ontogenetic variation. The transverse 
median swelling in the depression of small specimens 
(Fig. 2e, j) retains a faint expression in large cranidia. 
No bimodality can be detected in the strength of the 
librigenal ridge (Figs. 2n, o, 3b, c), which is 
consistently pronounced. 

In assigning this species to Proetus, 
Chatterton (1971) acknowledged its distance from the 
type species, the Wenlock P. concinnus (Dalman). 
However, several other Australian Emsian and Eifelian 
Proetinae are validly assigned to that genus. These 
include Proetus talenti Chatterton, 1971 (type of 
Devonoproetus Liitke, 1990), P. sparsinodosus Feist 
and Talent, 2000, and P. latimargo Feist and Talent, 
2000, the latter two originally assigned to 



Devonoproetus at the subgeneric level. Devonoproetus 
is a junior synonym of Proetus s.s. (Adrain 1997; Zhou 
et al. 2000). 

Proetus nemus was reassigned, with question, 
to the otherwise Ludlow-Lochkovian Ryckholtia 
Snajdr, 1980 (type Proetus ryckholti Barrande, 1846) 
by Liitke (1990). The new material described herein 
conflicts with this reassignment. Membership in 
Ryckholtia is precluded by the pronounced tuberculate 
sculpture on the glabella and axial rings of P. nemus, 
the strongly defined lateral occipital lobes, and sagittal 
elimination of the preglabellar field. 

This species displays characters that suggest 
alternative assignments. The elongate, tapering 
glabella of Proetus nemus and its pattern of sculpture 
(strong tuberculation posteriorly, becoming subdued 
anteriorly), together with the profile of the preglabellar 
region, including the wide (sag., exsag.) anterior 
cranidial border furrow, and the divergence of the 
facial suture between y and P resemble Longiproetus 
tenuimargo (Richter, 1909) (type of Longiproetus 
Cavet and Pillet, 1958). Longiproetus has been 
regarded as a synonym of Gerastos Goldfuss, 1843 
(Owens 1973), a valid subgenus of Gerastos (Snajdr 
1980), restricted to its type species on the basis of a 
distinctive shape of the rostral plate (Liitke 1990), or 
slightly expanded to include a small group of 
Rhenohercynian mid Eifelian to early Givetian species 
(Basse 1996, 2002). Liitke (1990) reassigned the 
Bohemian species that had been referred to 
Longiproetus (e.g., Snajdr 1980) to Coniproetus 
Alberti, 1966, and other genera, whilst the inadequately 
known Emsian species referred to Longiproetus by 
Pillet (1972) defy classification. Despite the similarities 
in the glabella and preglabellar region, several 
characters conflict with an alliance between P. nemus 
and Longiproetus. Notably, the strong genal ridge of 
P. nemus is lacking in L. tenuimargo and other certain 
congeners {sensu Basse 2002), the prominent lateral 
occipital lobes contrast with the inconspicuous lobes 
in Longiproetus s.s., L0 is wider than the basal part of 
the glabella, the cephalon is much less vaulted, the 
palpebral lobe is situated more posteriorly, and the 
pygidium is relatively paucisegmented (7+1 rings 
versus 8+1). The course of well preserved connective 
sutures on librigenae suggests that the rostral plate of 
P. nemus is more regularly trapezoidal or triangular 
than is that of L. tenuimargo (Liitke 1990:Text-fig. 8). 

Affinities to species that have been assigned 
to Devonoproetus by recent workers better account for 
the large occipital lobes, width of L0 relative to the 
glabella, and 7+1 pygidial segmentation. Among these, 
Proetus latimargo Feist and Talent, 2000 (Eifelian, 
Queensland) and P. zhusilengensis Zhou et al., 2000 



Proc. Linn. Soc. N.S.W., 125, 2004 



183 



EARLY DEVONIAN TRILOBITES FROM N.S.W. 



(Emsian, Inner Mongolia) resemble P. nemus in having 
a tongue-shaped glabella (narrowest in P. nemus) with 
dense, pronounced tuberculation, and P. latimargo 
shares the divergence of the facial suture between a 
and p. 

Among those species that have been referred 
to Devonoproetus, the strong genal ridge of Proetus 
nemus is developed in a group recognised by Basse 
(2002) as a separate genus, Rhenocynproetus, from 
which the Australian "Devonoproetus" species were 
explicitly excluded. The presence of a genal ridge in 
other genera of Proetinae [e.g. Gerastos: Snajdr 
1980:P1. 3, Fig. 13, PL 4, Fig. 17; Coniproetus 
(Bohemiproetus): Snajdr 1980:P1. 6, Figs 5, 6, 14; 
Lieberman 1994:Fig. 9.3) demonstrates that this feature 
is not an infallible indicator of relationships. Characters 
cited by Basse (2002) as excluding Australian species 
of Proetus from Rhenocynproetus also distinguish P. 
nemus; these include the large size of the lateral 
occipital lobes and weaker outer edge of the eye socle. 
Proetus nemus possesses (plesiomorphic) features 
considered by Basse (2002) to more generally 
distingish Proetus from Rhenocynproetus, such as a 
less inflated glabella, the lateral occipital lobes wider 
than the base of the glabella, terrace lines developed 
on the dorsal as well as lateral extent of the cranidial 
border, and the well developed librigenal spine. The 
presence of a pair of posterior border spines on the 
hypostome (Fig. 3d) is shared with Proetus (e.g. 
Whittington and Campbell 1967:P1. l,Fig. 17;Schrank 
1972:P1. 4, Fig. 7), including P. talenti, but is likely 
symplesiomorphic (Adrain 1997). 

Order CORYNEXOCHIDA Kobayashi, 1935 

Suborder SCUTELLUINA Hupe, 1953 

Family STYGINIDAE Vogdes, 1890 

Genus DENTALOSCUTELLUM Chatterton, 1971 

Type species 

Dentaloscutellum hudsoni Chatterton, 1971; by 
original designation. 

Dentaloscutellum hudsoni Chatterton, 1971 
Fig. 4a-i 

Dentaloscutellum hudsoni Chatterton, 1971:12-22, 
PI. 1, Figs 1-24, PI. 2, Figs 1-24, PI. 3, Figs 1-12, PI. 
24, Fig. 15, Text-figs 4-5. 

Material 

4 cranidia, 29 librigenae, 1 hypostome, 1 thoracic 
segment, 5 fragmentary pygidial margins. 



Discussion 

This species was fully described based on specimens 
from the Receptaculites Limestone near Taemas 
(Chatterton 1971). The Brogans Creek material is 
considered to be conspecific, the only possible 
difference being slightly more numerous cranidial 
tubercles (Fig. 4b, c) than in the type material. 

Order LICHIDA Moore, 1959 

Family LICHIDAE Hawle and Corda, 1847 

Subfamily TROCHURINAE Phleger, 1936 

Genus ACANTHOPYGE Hawle and Corda, 1847 

Type species 

Acanthopyge leuchtenbergii Hawle and Corda, 
1847; by subsequent designation of Reed (1902). 

Subgenus JASPERIA Thomas and Holloway, 1988 

Type species 

Acanthopyge (Mephiarges) bifida Edgell, 1955; by 
original designation. 

Acanthopyge (Jasperia) bifida Edgell, 1955 
Fig. 4j-t 

Acanthopyge (Mephiarges) bifida Edgell, 1955:138; 
Chatterton, 1971:30-41, PL 6, Figs 1-24, PL 7, Figs 
1-27, PL 8, Figs 1-17, Text-figs 8-10. 

Material 

7 cranidia, 1 rostral plate, 7 librigenae, 3 
hypostomes, 1 thoracic segment, 2 pygidia. 

Discussion 

The Brogans Creek specimens are indistinguishable 
from those described from Wee Jasper (Edgell 1955) 
and Taemas (Chatterton 1971). The species was fully 
described by Chatterton (1971), rendering description 
of the Brogans Creek material unnecessary. A few 
specimens are illustrated (Fig. 4j-t) in support of the 
conspecificity of the collections. 

Order PHACOPIDA Salter, 1864 

Suborder CALYMENTNA Swinnerton, 1915 

Family CALYMENIDAE Milne Edwards, 1840 

Genus STHENAROCALYMENE Siveter, 1977 

Type species 

Sthenarocalymene lirella Siveter, 1977; by original 
designation. 



184 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE AND A.J. WRIGHT 




Figure 4. a-i, Dentaloscutellum hudsoni Chatterton, 1971. Scale bars 1mm. a, AMF 124725, librigena, 
dorsal view; b-d, AMF 124726, cranidium, dorsal, anterior and lateral views; e, AMF 124727, librigena, 
dorsal view; f, AMF 124728, librigena, ventral view; g, AMF 124729, fixigena, dorsal view; h, AMF 
124730, incomplete pygidium, ventral view; i, AMF 124731, incomplete pygidium, ventral view, j-t, 
Acanthopyge (Jasperia) bifida Edgell, 1955. Scale bars 1 mm. j, AMF 124732, rostral plate, ventral view; 
k, AMF 124733, cranidium, dorsal view; 1-m, AMF 124734, cranidium, dorsal and anterior views; n, 
AMF 124735, librigena, dorsal view; o-q, AMF 124736, pygidium, lateral, dorsal and ventral views; r, 
AMF 124737, librigena, ventral view; s-t, AMF 124738, hypostome, ventral and dorsal views. 



Proc. Linn. Soc. N.S.W., 125, 2004 



185 



EARLY DEVONIAN TRILOBITES FROM N.S.W. 



Sthenarocalymene sp. 

Material 

Two cranidial fragments, one fragmentary librigena. 

Discussion 

A few calymenid cephalic fragments indicate the 
presence of a species lacking a buttress between the 
fixigena and L2. On this basis the material is assigned 
to Sthenarocalymene, the non-buttressed calymenid 
in many Australian Lower Devonian faunas [see 
Sandford (2000) for discussion of this genus, its 
synonym Apocalymene Chatterton and Campbell, 
1980, and Gravicalymene Shirley, 1936]. The Brogans 
Creek material may be identical with S. quadrilobata 
(Chatterton, 1 97 1 ), which co-occurs with the other taxa 
described herein in the lower Receptaculites Limestone 
at Locality T of Chatterton (1971), but specific identity 
requires better specimens. 



ACKNOWLEDGEMENTS 

Alex Cook (Queensland Museum) assisted AJW 
in the field, and processed and picked much of the material 
studied here. Yongyi Zhen (Australian Museum) 
photographed the specimens and assembled the plates, and 
processed several blocks of limestone. Robert Owens 
(National Museum of Wales) provided helpful suggestions 
on an earlier version of the mansucript. 



REFERENCES 

Adrain, J.M. (1997). Proetid trilobites from the Silurian 
( Wenlock-Ludlow) of the Cape Phillips 
Formation, Canadian Arctic Archipelago. 
Palaeontographia Italica 84, 21-111. 

Alberti, G.K.B. (1966). Uber einige neue Trilobiten aus 
dem Silurium und Devon, besonders von 
Marokko. Senckenbergiana lethaea 47, 111- 
121. 

Barrande, J. (1846). 'Notice preliminaire sur le systeme 
Silurien et les Trilobites de Boheme'. (Leipzig). 

Basden, A., Burrow, C, Hocking, M., Parkes, R. and 
Young, G. (2002). Siluro-Devonian 
microvertebrates from southeastern Australia. 
Courier Forschungsinstitut Senckenberg 223, 
201-222. 

Basse, M. (1996). Trilobiten aus mittelerem Devon des 
Rhenohercynikums: I. Corynexochida und 
Proetida (1). Palaeontographica A 239, 89-182. 

Basse, M. (2002). 'Eifel-Trilobiten. 1. Proetida'. 
(Goldschneck-Verlag: Korb). 

Campbell, K.S.W. and Talent, J.A. (1963). 

Malurostrophia, a new genus of stropheodontid 
brachiopod from the Devonian of Australia. 



Proceedings of the Royal Society of Victoria 80, 

309-330. 
Carne, J.E. and Jones, L.J. (1919). The limestone deposits 

of New South Wales. Geological Survey of New 

South Wales, Mineral Resources 25, 1-411. 
Cavet, P. and Pillet, J. (1958). Les trilobites du calcaires a 

Polypiers silicieux (Eifelien) du Synclinal de 

Villefranche-de-Confluent (Pyrenees 

Orientales). Bulletin de la Societe geologique de 

France 81, 21-37. 
Chatterton, B.D.E. (1971). Taxonomy and ontogeny of 

Siluro-Devonian trilobites from near Yass, New 

South Wales. Palaeontographica A 137, 1-108. 
Chatterton, B.D.E. (1973). Brachiopods of the 

Murrumbidgee Group, Taemas, New South 

Wales. Bulletin of the Bureau of Mineral 

Resources, Geology and Geophysics (Australia) 

137, 1-146. 
Chatterton, B.D.E. and Campbell, K.S.W. (1980). Silurian 

trilobites from near Canberra and some related 

forms from the Yass Basin. Palaeontographica 

A 167, 77-119. 
Colquhoun, G.P. (1995). Early Devonian conodonts from 

the Capertee High, E Lachlan Fold Belt, 

southeastern Australia. Courier 

Forschungsinstitut Senckenberg 182, 347-369. 
Colquhoun, G.P. (1998). Early Devonian stratigraphy, 

conodont faunas and palaeogeography of the 

Capertee High. PhD thesis, University of 

Wollongong. 
Colquhoun, G.P., Meakin, N.S. and Watkins, J.J. (1999). 

Kandos Group. In 'Dubbo 1:250 000 Geological 

Sheet, 2 nd edition. Explanatory Notes' (Eds N.S. 

Meakin and E.J. Morgan) pp. 125-143. 

(Geological Survey of New South Wales: 

Sydney). 
Dalman, J.W. (1827). Om Palaeaderna eller de sa kallade 

Trilobiterna. Kungliga Svenska 

Vetenskapsakademiens Handlingar 1826, 113- 

152, 226-294. 
Edgell, H.S. (1955). A Middle Devonian lichid trilobite 

from South-Eastern Australia. Palaontologische 

Zeitschrift 29, 136-145. 
Feist, R. and Talent, J.A. (2000). Devonian trilobites from 

the Broken River region of northeastern 

Australia. Records of the Western Australian 

Museum, Supplement 58, 65-80. 
Fortey, R.A. and Owens, R.M. (1975). Proetida - a new 

order of trilobites. Fossils and Strata 4, 227- 

239. 
Garratt, M.J. and Wright, A.J. (1988). Late Silurian to 

Early Devonian biostratigraphy of southeastern 

Australia. In 'Devonian of the World. Volume 

HI: Paleontology, Paleoecology and 

Biostratigraphy' (Eds N.J. McMillan, AF. 

Embry, Jr., and D.J. Glass) pp. 647-662. 

(Canadian Society of Petroleum Geologists: 

Calgary). 
Goldfuss, A. (1843). Systematische Ubersicht der 

Trilobiten und Beschreibung einiger neuen Arten 



186 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE AND A.J. WRIGHT 



derselben. Neues Jahrbuch fur Mineralogie, 
Geognosie, Geologie und Petrefaktenkunde 1843, 
537-567. 

Hawle, I. and Corda, A.J.C. (1847). Prodrom einer 
Monographic der bohmischen Trilobiten. 
Bohmischen Gesellschaft der Wissenschqften, 
Abhandlungen 5, 1-176. 

Hupe, P. (1953). Classe des Trilobites. In Traite de 

Paleontologie. Volume 3' (Ed J. Piveteau) pp. 
44-246. (Masson: Paris). 

Kobayashi, T. (1935). The Cambro-Ordovician Formations 
and Faunas of South Chosen. Palaeontology. Part 
HI. Journal of the Faculty of Science, Tokyo 
University 4, 49-344. 

Lieberman, B.S. (1994). Evolution of the trilobite 

subfamily Proetinae Salter, 1864, and the origin, 
diversification, evolutionary affinity, and 
extinction of the Middle Devonian proetid fauna 
of eastern North America. Bulletin of the 
American Museum of Natural History 223, 1- 
176. 

Lindley, I.D. (2002). Acanthodian, onychodontid and 

osteolepid fish from the middle-upper Taemas 
Limestone (Early Devonian), Lake Burrinjuck, 
New South Wales. Alcheringa 26, 103-126. 

Lishmund, S.R., Dawood, A.D. and Langley, W.V. 

(1986). The limestone deposits of New South 
Wales, 2 nd edition. Geological Survey of New 
South Wales, Mineral Resources 25, 1-373. 

Liitke, F. (1990). Contributions to a phylogenetical 
classification of the subfamily Proetinae 
SALTER, 1864 (Trilobita). Senckenbergiana 
lethaea 71, 1-83. 

Mawson, R. (1987). Early Devonian conodont faunas 

from Buchan and Bindi, Victoria. Palaeontology 
30, 251-297. 

Meakin, N.S. and Morgan, E.J. (1999). 'Dubbo 1:250 000 
Geological Sheet, 2 nd edition. Explanatory 
Notes'. (Geological Survey of New South 
Wales: Sydney). 

Milne Edwards, H. (1840). 'Histoire Naturelle des 
Crustaces, Comprenant l'Anatomie, la 
Physiologie et la Classification de ces Animaux, 
3'. (Roret: Paris). 

Moore, R.C. (1959). Order Lichida. In 'Treatise on 

Invertebrate Paleontology. Part O. Arthropoda 
1' (Ed R.C. Moore) p. 0495. (Geological 
Society of America, University of Kansas Press: 
Lawrence). 

Owens, R.M. (1973). British Ordovician and Silurian 
Proetidae (Trilobita). Palaeontographical 
Society Monograph, 1-98. 

Pedder, A.E.H., Jackson, J.H. and Philip, G.M. (1970). 
Lower Devonian biostratigraphy in the Wee 
Jasper region of New South Wales. Journal of 
Paleontology 44, 206-251. 

Phleger, F.B. (1936). Lichadian trilobites. Journal of 
Paleontology 10, 593-615. 

Pickett, J.W. (1972). Correlation of the Middle Devonian 
formations of Australia. Geological Society of 



Australia 18, 457-466. 
Pickett, J.W. (1978). Conodont faunas from the Mount 

Frame Limestone (Emsian-Eifelian). Bulletin of 

the Bureau of Mineral Resources, Geology and 

Geophysics (Australia) 192, 97-107. 
Pillet, J. (1972). Les trilobites du Devonien inferieur et du 

Devonien moyen du Sud-Est du Massif 

Armoricain. Memoires Societe d' Etudes 

scientifiques de I'Anjou 1, 1-307. 
Reed, F.R.C. (1902). Notes on the genus Lichas. Journal 

of the Geological Society of London 58, 59-82. 
Richter, R. (1909). Beitrage zur Kenntnis devonischer 

Trilobiten aus dem Rheinischen Schiefergebirge. 

Vorbericht zu einer Monographic der Trilobiten 

der Eifel. Dissertation, Marburg. 
Salter, J.W. (1864). A monograph of the British trilobites 

from the Cambrian, Silurian, and Devonian 

formations. Monographs of the 

Palaeontographical Society, 1-80. 
Sandford, A.C. (2000). Trilobite faunas and 

palaeoenvironmental setting of the Silurian 

(early Ludlow) Melbourne Formation, central 

Victoria. Alcheringa 24, 153-206. 
Schrank, E. (1972). Proetacea, Encrinuridae und 

Phacopina (Trilobita) aus silurischen 

Geschieben. Geologie 76, 1-117. 
Shirley, J. (1936). Some British trilobites of the family 

Calymenidae. Quarterly Journal of the 

Geological Society of London 92, 384-421. 
Siveter, D.J. (1977). The Middle Ordovician of the Oslo 

region, Norway, 27. Trilobites of the family 

Calymenidae. Norsk Geologisk Tidsskrift 56, 

335-396. 
Snajdr, M. (1980). Bohemian Silurian and Devonian 

Proetidae (Trilobita). Rozpravy Ustredniho 

ustavu geologickeho 45, 1-324. 
Steininger, J. (1831). 'Bemerkungen iiber die 

Versteinerungen, welche in dem 

UbergangsKalkgebirge der Eifel gefunden 

, werden' . (Eine Beilage zum Gymnasial-Program 

zu Trier: Trier). 
Swinnerton H.H. (1915). Suggestions for a revised 

classification of trilobites. Geological Magazine, 

New Series 2, 407-496, 538-545. 
Talent, J.A., Mawson, R. and Simpson, A.J. (2000). 

Silurian to Early Carboniferous (Tournaisian) 

platform-slope sequences in eastern Australia: 

recent advances in stratigraphic alignments. 

Geological Society of Australia, Abstracts 61, 

114-120. 
Thomas, AT. and Holloway, D.J. (1988). Classification 

and phylogeny of the trilobite order Lichida. 

Philosophical Transactions of the Royal Society 

of London 32, 179-262. 
Vogdes, A.W. (1890). A bibliography of Paleozoic 

Crustacea from 1698 to 1890 including a list of 

North American species and a systematic 

arrangement of genera. Bulletin of the United 

States Geological Survey 63, 1-177. 



Proc. Linn. Soc. N.S.W., 125, 2004 



187 



EARLY DEVONIAN TRILOBITES FROM N.S.W. 



Whittington H.B. and Campbell, K.S.W. (1967). Silicified 
Silurian trilobites from Maine. Bulletin of the 
Museum of Comparative Zoology 135, 447-483. 

Zhou Z.-q., Siveter, D.J. and Owens, R.M. (2000). 
Devonian proetid trilobites from Inner 
Mongolia. Senckenbergiana lethaea 79, 459- 
499. 



Proc. Linn. Soc. N.S.W., 125, 2004 



A New Species of the Henicopid Centipede Dichelobius 
(Chilopoda: Lithobiomorpha) from Southeastern Australia and 

Lord Howe Island 

Gregory D. Edgecombe 

Australian Museum, 6 College Street, Sydney, NSW 2010 
(greged @ austmus .gov . au) 



Edgecombe, G.D. (2004) A new species of the henicopid centipede Dichelobius (Chilopoda: Lithobiomorpha) 
from southeastern Australia and Lord Howe Island. Proceedings of the Linnean Society of New South 
Wales 125, 189-203. 

The genus Dichelobius Attems, 191 1, based on D. flavens Attems, 1911, from the southwest of Western 
Australia, has its only other previously assigned species in New Caledonia and Chile. The Tasmanian type 
species of the monotypic Tasmanobius Chamberlin, 1920, is regarded as a member of Dichelobius. 
Dichelobius giribeti n. sp. represents the genus in eastern mainland Australia (southeastern New South 
Wales, the Australian Capital Territory, and northeastern Victoria) and on Lord Howe Island. Dichelobius 
bicuspis Ribaut, 1923, is widely distributed in New Caledonia. 

Manuscript received 1 March 2003, accepted for publication 8 January 2004. 

KEYWORDS: Anopsobiinae, Chilopoda, Dichelobius, Henicopidae, Lithobiomorpha. 



INTRODUCTION 

The subfamily Anopsobiinae is a group of 
minute centipedes (Chilopoda) in the predominantly 
southern temperate family Henicopidae. Anopsobiinae 
is distributed chiefly in the Southern Hemisphere, with 
species described from Patagonian Argentina and Chile 
(Silvestri 1899, 1909a-b; Verhoeff 1939; Chamberlin 
1962), the Falkland Islands (Eason 1993), New 
Zealand (Silvestri 1909a; Archey 1917, 1937), New 
Caledonia (Ribaut 1923), Tasmania (Chamberlin 
1920), New South Wales (Edgecombe 2003), 
southwest Western Australia (Attems 1911), and the 
Cape region of South Africa (Attems 1928). Four 
Gondwanan genera have been named: Anopsobius 
Silvestri, 1899, Catanopsobius Silvestri, 1909b, 
Dichelobius Attems, 1911, and Tasmanobius 
Chamberlin, 1920. Four additional anopsobiine genera, 
all monotypic, occur in the Northern Hemisphere, 
namely Anopsobiella Attems, 1938, Ghilaroviella 
Zalesskaja, 1975, Shikokuobius Shinohara, 1982, and 
Rhodobius Silvestri, 1933. In total, 17 species and 
subspecies of Anopsobiinae have been described. 

Silvestri (1909a) cited the occurrence of an 
anopsobiine from Sydney, but formal descriptions of 
Anopsobiinae in eastern Australia are limited to 
Tasmanobius relictus Chamberlin, 1920, based upon 
a single specimen from Tasmania, and Anopsobius 
wrighti Edgecombe, 2003, from northern New South 



Wales. Mesibov (1986) indicated the presence of two 
species of Anopsobiinae in Tasmania. The present 
study continues a systematic treatment of 
Anopsobiinae of Australia by documenting a new 
species of Dichelobius from New South Wales, the 
Australian Capital Territory, Victoria, and Lord Howe 
Island (Fig. 1). 

For electron microscopy, specimens were 
photographed on a Leo 435VP using a Robinson 
backscatter detector. Digital images were assembled 
into plates with Photoshop. Morphological 
terminology is as summarised by Edgecombe 
(2001:203), with terminology for the mandible as in 
Edgecombe et al. (2002:40, Fig. 4). 

The following abbreviations are used for 
repositories of specimens examined: 
AM - Australian Museum, Sydney 
ANIC - Australian National Insect Collection, 
Canberra 

MCZ - Museum of Comparative Zoology, Harvard 
University, Cambridge, MA 
MNHN - Museum National d'Histoire Naturelle, 
Paris 

NMW - Naturhistorisches Museum Wien 
QM - Queensland Museum, Brisbane 
WAM - Western Australian Museum, Perth. 
Other abbreviations: Berl., ANIC Berlesate; CBCR, 
Australian Museum Centre for Biodiversity and 
Conservation Research; Ck, Creek; Mt, Mountain; NP, 
National Park; rf, rainforest; SF, State Forest. 



A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 



Brisbane 



Lord Howe 
Island © 




Figure 1. a, southeastern Australia and Lord Howe 
Island. Inset shows location of map in b, indicating 
records of Dichelobius giribeti n. sp. (open dots) in 
New South Wales, the Australian Capital Territory, 
and Victoria. 

Collectors: GBM - G.B. Monteith; JFL - J.F. 
Lawrence; RJB - R.J. Brooks; RWT - R.W. Taylor. 

Order LITHOBIOMORPHA Pocock, 1902 

Family HENICOPIDAE Pocock, 1901 

Subfamily ANOPSOBIINAE Verhoeff, 1907 

Genus DICHELOBIUS Attems, 1911 

Tasmanobius Chamberlin, 1920 n. syn. 

Type species 

Dichelobius flavens Attems, 1911; by 
monotypy. 



Assigned species 

Dichelobius relictus (Chamberlin, 1920) n. 
comb.; D. bicuspis Ribaut, 1923; D. schwabei 
Verhoeff, 1939; D. giribeti n. sp. 

Diagnosis 

Anopsobiinae with spiracle on segments 3, 
10 and 12, variably present on segment 14. 

Discussion 

The Gondwanan genera Dichelobius, 
Tasmanobius and Anopsobius share several 
apomorphic characters relative to Northern 
Hemisphere Anopsobiinae. These include coxal pores 
confined to legs 14 and 15, a ventrodistal spur on the 
prefemur of legs 14 and 15, an elongate longitudinal 
median furrow on the head shield, the basal article of 
the female gonopod extended as a short process bearing 
the spurs, and indistinct scutes on the proximodorsal 
part of the pretarsal claws (Edgecombe and Giribet 
2003). Considering previous concepts of Dichelobius 
(Attems 1928; Verhoeff 1939; Shinohara 1982), 
reduced spiracles are the only morphological character 
that unites its members to the exclusion of Anopsobius 
as delimited by Chamberlin (1962) and Edgecombe 
(2003). The Dichelobius distribution of spiracles is 
shared by the eastern Australian species D. giribeti. 
The cladistic reliability of a diminished number of 
segments with spiracles can be questioned because 
other genera of Anopsobiinae have been diagnosed 
based on having spiracles confined to segments 3, 10 
and 12 {Tasmanobius), 3, 12 and 14 (Rhodobius) or 3 
and 10 only (Catanopsobius). However, molecular 
sequence data provide independent support for a close 
relationship between D. flavens and D. giribeti, with 
the implication that their shared spiracle distribution 
can be considered a synapomorphy (Fig. 2a). 
Parsimony analysis of five molecular loci as well as 
combination of the molecular data and morphology 
unite D. flavens and D. giribeti to the exclusion of 
Anopsobius species under many explored gap costs 
and transversiomtransition ratios (Edgecombe and 
Giribet 2003) (Fig. 2c). An alternative relationship 
between D. giribeti and Anopsobius (Fig. 2b) is 
discussed below. 

Verhoeff (1925) cited the presence of a 
median suture in the maxillipede pleural band as an 
additional character by which Dichelobius is 
distinguished from Anopsobius. The presence of a 
median suture (see Fig. 6j) is a plesiomorphic character, 
shared with Henicopinae, and is thus not useful for 
defining Dichelobius as a clade. 

Tasmanobius relictus Chamberlin, 1920, is 
considered to be a member of Dichelobius as grouped 



190 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE 



s 

| 

•c 



§ 

""§3 

8 5 

St 



CO 

o 
u 

a 

(0 

CD 



3 

■9 

Q 5: 



.s 

■5 
fife 

■c -o 

Q 8) 



V) 

3 

•5 
o 
</> 

Q. 
o 

c 



3 

5 
o 

I 

e 



to 

•Q £ 

O (B 



I I 



ir 1 



.3 
■5 

0) c 

■c 



.3 

■5 

a> a> 

■c .-Q 



Q 55 Q O) 



(0 

J* 

.3 

o 
</> 

& 

c 



i 

3 

■i 

c 



(0 

•2 .a 

•9 5 



T 



2 4 

Tv : Ts cost 



w 



0) 



aw 

S o 
^ c 



I I 






inf. 



Figure 2. a, b, alternative cladograms for 
Anopsobiinae based on combined morphological 
and molecular data (Edgecombe and Giribet 2003). 
Character 1, absence of spiracles on segment 8; 
character 2, short posteroventral spine on pretarsal 
claw; c, summary of 12 analyses for combined 
morphological and molecular data with different 
gap costs (gap: substitution = 1:1, 2:1, 4:1) and 
transversion: transition costs (1:1, 2:1, 4:1, infinity). 
Black squares, parameters that resolve cladogram 
a (Dichelobius monophyletic); white squares, 
parameters that resolve cladogram b (Dichelobius 
paraphyletic); grey square, cladograms a and b of 
equal length. 



herein (with Tasmanobius consequently being a junior 
subjective synonym of Dichelobius). Tasmanobius 
relictus was described as having spiracles on segments 
3, 10, and 12, as in Dichelobius. Mesibov (1986) 
suggested that a widespread Tasmanian anopsobiine 
species (Anopsobiine sp. 2 of Mesibov 1986) may be 
Tasmanobius relictus, and that species closely 
resembles Dichelobius giribeti. The holotype and sole 
type specimen of T. relictus (MCZ 14533) is in poor 
condition, and lacks locality data more specific than 
Tasmania, making the identification of any other 
specimen as this species problematical. The description 
by Chamberlin did not note a spiracle on segment 14 
which is present in the Tasmanian Dichelobius, though 
this is not obvious in contracted specimens, as noted 
by Mesibov (1986). A spiracle being absent on segment 
eight in T. relictus and the colour being "nearly 
chestnut" (Chamberlin 1920) make it probable that this 
species is identical with the Tasmanian Dichelobius 
(=Anopsobiinae sp. 2 of Mesibov 1986) rather than 
the northwestern Tasmanian Anopsobius 
(= Anopsobiinae sp. 1 of Mesibov 1986), which has a 
spiracle on segment 8 and is more orange-yellow than 
orange-brown. Accordingly, the name Dichelobius 
relictus (Chamberlin, 1920) is applied to Anopsobiinae 
sp. 2 of Mesibov (1986). 

Attems' (1928:74) key to anopsobiine genera 
followed Chamberlin' s (1920) in distinguishing 
Dichelobius and Tasmanobius based on the former 
having a 1-jointed tarsus 13 and the latter a 2-jointed 
tarsus 13. This distinction is inconsistent with the 
referral of D. bicuspis, which has a 2-jointed tarsus 13 
(even fide Attems 1928:77). The supposed difference 
between these species seems to be nothing more than 
a terminological difference in what constitutes a 
"joint", since D. flavens, D. bicuspis and D. relictus 
are, upon direct comparison, identical with respect to 
the segmentation of leg 13. All have a distinct 
articulation on the tarsus of leg 13, though it is less 
flexed than is the articulation on leg 14. 

Other ambiguities concerning Attems' 
description and illustrations of Dichelobius flavens 
have plagued previous interpretations of the genus, and 
exaggerated differences between D. flavens and other 
species. Interpretation of D. flavens is based on 
examination of syntypes from Lion Mill (WAM), 
Freemantle and Eradu (NMW), and large new 
collections from the southwest of Western Australia 
(AM, ANIC, WAM). Dichelobius bicuspis and D. 
schwabei were distinguished from D. flavens by the 
first two species having two coxal pores on legs 14 
and 15 in the female, versus a single pore on each of 
the coxae in D. flavens. This cannot be upheld, since 
large females of D. flavens characteristically have two 



Proc. Linn. Soc. N.S.W., 125, 2004 



191 



A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 



coxal pores on both legs 14 and 15. Attems' (1911:157, 
Fig. 10) described and figured a single spur on the 
female gonopod in D. flavens, which Ribaut (1923) 
and Verhoeff (1939) cited as a distinction from the 
pair of spurs in D. bicuspis and D. schwabei, 
respectively. Large specimens of Dichelobius flavens 
resemble congeners (and indeed all other 
Anopsobiinae) in having a pair of spurs. The specimen 
drawn by Attems, with a single spur and single coxal 



pore, is typical of immature stadia of all Dichelobius 
species (see Archey 1937:pl. 23, fig. 6, for a 
comparable stage in Anopsobius neozelanicus). Ribaut 
(1923:27) distinguished D. bicuspis by its plumose 
setae along the length of the inner margin of the distal 
article of the telopodite of the first maxilla versus only 
three plumose setae confined to the distal end of this 
article in D. flavens (Attems 1911:Fig. 3). Either 
Attems' drawing is erroneous or else the illustrated 




Figure 3. Pretarsal claws in Anopsobiinae. a, Dichelobius relictus (Chamberlin, 1920). Leg 14, posterior 
side, b, c, Dichelobius flavens Attems, 1911. Leg 14, posterior and anterior sides, d, Anopsobius neozelanicus 
Silvestri, 1909. Leg 14, posterior side, e, f, Shikokuobius japonicus (Murakami, 1967). Leg 13, posterior 
and anterior sides. Scales 10 \xm except b, 5 u\m. 



192 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE 



specimen is anomalous, because D. flavens has 
plumose setae all along the inner margin of this article, 
the same d&D. bicuspis (Ribaut 1923:Figs. 30, 31) and 
other congeners. 

Certain characters of the pretarsus (claws) 
conflict with the monophyly of Dichelobius as grouped 
herein. Dichelobius flavens (Fig. 3b, c) andD. bicuspis 
differ from D. giribeti (Fig. 8b) and D. relictus (Fig. 
3a) in having a long, needle-like spine (=" sensory spur" 
of Eason 1964:Fig.486) originating ventrally on the 
posterior side of the main claw. In the latter two species, 
the posteroventral spine is short, and a short spine is 
shared by species of Anopsobius, such as A. 
neozelanicus Silvestri, 1909a (Fig. 3d) and A. wrighti 
(Edgecombe 2003:Figs.30, 31). The short spine 
appears to be apomorphic within the Gondwanan group 
of Anopsobiinae (i.e., a clade composed of Anopsobius 
+ Dichelobius) because the Japanese anopsobiine 
Shikokuobius japonicus resembles Dichelobius flavens 
and D. bicuspis in possessing a greatly elongated 
posteroventral spine (Fig. 3e, f). The cladogram 
implied by this character, in which D. giribeti is more 
closely related to Anopsobius than to D. flavens (Fig. 
2b), is retrieved under several parameter sets for 
combined morphological and molecular data (Fig. 2c). 
This cladogram would favour the assignment of D. 
giribeti to another genus. Should this topology find 
further support from additional data, Tasmanobius 
Chamberlin, 1920, could be rediagnosed to receive D. 
giribeti. A rediagnosed concept of that genus might 
emphasise the shared 14-15 antennal articles, short 
pretarsal posteroventral spine, absence of a distal 
spinose projection on the tibia of leg 12, and lack of 
spiracles on segments 5 and 8. 

Key to Dichelobius species 



4a. Tibia of leg 12 with short, blunt distal 

projection flavens Attems, 1911 [Western 

Australia] 

4b. Tibia of leg 12 with spinose distal 

projection bicuspis Ribaut, 1923 [New 

Caledonia] 

Dichelobius giribeti n. sp. 

Dichelobius sp. Edgecombe, 2004:Fig. 38A. 
Dichelobius sp. ACT. Edgecombe and Giribet, 
2003:Figs. 1-3. 

Etymology 

For Gonzalo Giribet, my collaborator in 
henicopid phylogeny, who sequenced DNA from this 
species. 

Diagnosis 

Dichelobius usually with 15 antennal articles; 
head pale orange, tergites orange-yellow; four to six 
(most commonly five) teeth on each dental margin of 
maxillipede; spiracle lacking on segment 14; two coxal 
pores on legs 14 and 15 in females, one or two pores 
on both legs in males; short posteroventral spine on 
pretarsus. 

Type material 

Holotype: AM KS 82628, female (Fig. 4b), 
Badja SF, NSW, Peters Rd, 36°08'52"S 149°32'09"E, 
J. Tarnawski and S. Lassau, 13.iii. 1999; length of body 
5.1 mm. Paratypes, all from type locality, same 
collection: AM KS 82629, male (Fig. 4c), KS 82630, 
male (Fig. 5b-e), KS 82631, female (Figs. 6a-g, 7a, b, 
d, h, j-1, 8k), KS 82632, female (Fig. 8i, j, n), KS 82633, 
male (Fig. 81), KS 82634, 10 females, 1 male. 



la. Dental margin of maxillipede coxosternite 

lacking median notch schwabei Verhoeff, 

1939 [Chile] 

lb. Dental margin of maxillipede coxosternite with 

median notch 2 

2a. 14-15 (usually 15) antennal articles; pretarsus 
with short posteroventral spine, not more than one- 
eighth length of main claw (Fig. 8b) 3 

2b. 17 antennal articles; pretarsus with needle-like 
posteroventral spine nearly as long as main claw 
(Fig. 3c) 4 

3a. Spiracle absent on segment 14 giribeti n. sp. 

[southeastern Australia, Lord Howe Island] 

3b. Spiracle present on segment 14 relictus 

Chamberlin, 1920 [Tasmania] 



Other material 

NSW: AM KS 82635, Kanangra-Boyd NP, 
Empress Fire Trail turnoff, 33°59'S 150°08'E, M. 
Gray, G. Hunt and J. McDougall, 27.iii.1976, 
Eucalyptus pauciflora; AM KS 82636, female (Figs 
4a, 5a), KS 82637, female (Fig. 8b, e), KS 82638, male 
(Fig. 6i, j), Monga SF, NSW, Link Rd, 35°34'04"S 
149°54' 14"E, R. Harris and H. Smith, 16.iii.1999; AM 
KS 82639, Buckenbowra SF, Macquarie Rd, 70 m S 
from junction with Milo Rd, 35°38' 15"S 149°53'27"E, 
1020 m, L. Wilkie and R. Harris, 16.iii.1999; AM KS 
82640, Tallaganda SF, South Forest Way, 35°42'50"S 
149°32'20"E, J. Tarnawski and S. Lassau, 15.iii.1999; 
AM KS 82641, Dampier SF, Coomerang Rd, 
36°04'01"S 149°54'57"E, R. Harris and H. Smith, 
11. hi. 1999; AM KS 82642, Badja SF, Wiola Ck Fire 
Trail, 36°05.56'S 149°35.09'E, J. Tarnawski and S. 
Lassau, 13.iii.1999; AM KS 82643, Badja SF, Burkes 



Proc. Linn. Soc. N.S.W., 125, 2004 



193 



A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 










Figure 4. a-c, Dichelobius giribeti n. sp. a, AM KS 82636, female, Monga SF, NSW. b, holotype AM KS 
82628, female, Badja SF, NSW, terminal segments and gonopods; c, AM KS 82629, male, Badja SF, 
NSW, terminal segments and gonopods. All scales 100 |im. 



194 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE 



Rd, 36°10'33"S 149°31'58"E, J. Tarnawski and S. 
Lassau, 13.iii.1999; AM KS 82644, Badja SF, Burkes 
Rd, approx. 1.3 km E from junction with Peters Rd, 
36°10.55'S 149°31.97'E, 992 m, J. Tarnawski and S. 
Lassau, 13.iii.1999; AM KS 82645, Bodalla SF, 300 
m along Reservoir Link Rd from junction with Big 
Rock Rd, 36°07.25'S 150°2.82'E, 121 m, L. Wilkie 
and R. Harris, 09.iii.1999; AM KS 82646, Bodalla SF, 
Orange Ridge Rd, 36°16'55"S 149°53'31"E, R. Harris 
and H. Smith, 12.nj.1999; AM KS 82647, Wadbilliga 
NP, 9.6 km N on Bumberry Ck Fire Trail, 36°14.33'S 
149°33.60'E, 1059 m, L. Wilkie and R. Harris, 
13.iii.1999. 

ANIC (ex. Berl. 855), Kanangra-Boyd NP, 
W Morong Creek, 33°58'S 150°04'E, 1200 m, L. Hill, 
03.X.1982; ANIC (ex. Berl. 829), Kanangra-Boyd NP, 
Kanangra Brook and Rocky Spur, 34°00'S 150°06'E, 
L. Hill, 20.iii. 1982, closed forest; ANIC (ex. Berl. 852) 
Twin Falls, 14 km SE Moss Vale, 34°39'S 150°28'E, 
600 m, L. Hill, ll.vii.1982; ANIC (ex. Berl. 663), 
Pigeon House Range via Nerriga, 35°02'S 150°08'E, 
J.C. Cardale, 22.xi.1979; ANIC (ex. Berls 2, 18, 34, 
78A, 206A, 222, 246, 468, 657, 851), Clyde Mt, 
35°33'S 149°57'E, 500-c. 800 m, various collections 
1966-1982, dry sclerophyll, wet sclerophyll, rf; ANIC 
(ex. Berl. 877), 2 km N Monga, 35°34'S 149°56'E, 
M.S. Harvey, 18.ix.1983, wet sclerophyll; ANIC (ex. 
Berl. 594), Monga, 35°35'S 149°55'E, JFL and T. 
Weir, 10.iii.1978, wet sclerophyll; ANIC (ex. Berl. 
739), Tallaganda SF, 7 km ENE Captains Flat, 35°34'S 
149°31'E, W. Allen, 29.viii.1981; ANIC (ex. Berl. 
1069), Kioloa SF, 35°35'S 150°18'E, JFL and N. 
Lawrence, 4-5.iii.1986; ANIC (ex. Berl. 927), Milo 
Forest Preserve, 1.6 km S Monga, 35°36'S 149°55'E, 
L. Hill, 25.xii.1983; ANIC (ex. Berl. 218), 8.8 kmESE 
Captains Flat, 35°38'S 149°31'E, 940 m, RWT, 
10.U970, dry sclerophyll; ANIC (ex. Berl. 891), 
Rosedale, 35°49'S 150°14'E, R.J. Moran, 20.xi.1983, 
eucalypt litter; ANIC (ex. Berl. 933), Kosciusko NP, 
1 km ENE Mt Sunrise, 36°22'S 148°29'E, L. Hill, 
4.ii.l984; ANIC (ex. Berl. 935), Kosciusko NP, 4 km 
NNEMtPerisher, 36°22'S 148°29'E, L. Hill, 4.ii.l984; 
ANIC (ex. Berl. 10), Brown Mt, 36°36'S 149°23'E, c. 
3000 ft., RWT, 5.L1967, wet sclerophyll; ANIC (ex. 
Berl. 20), Brown Mt, c. 2800 ft., RWT and R.J. Bartell, 
30.iii.1967, rf; ANIC (ex. Berl. 24), Brown Mt, 2500- 
3000 ft., RWT and R.J. Bartell, 1 l.iv.1967; ANIC (ex. 
Berl. 41), Brown Mt, Rutherford Creek, 2700 ft., RWT 
and RJB, 9.xii.l967, rf; ANIC (ex. Berl. 42), Brown 
Mt, c. 3000 ft., RWT and RJB, 9.xii.l967, rf. 

ACT: ANIC (ex. Berl. 283), Black Mt, 
eastern slope, 35°16'S 149°06'E 750 m, J. Simmons, 
26.V.1970, dry sclerophyll; ANIC (ex. Berl. 228), 
Uriarra to Piccadilly Circus, 35°19'S 148°51'E, 700 
m, RWT, 27.L1970, dry sclerophyll; ANIC (ex. Berl. 



225), Uriarra to Piccadilly Circus, 35°20'S 148°50'E, 
500 m, RWT, 16.L1970, wet sclerophyll; ANIC (ex. 
Berl. 231), Uriarra to Piccadilly Circus, 35°20'S 
148°50'E, 1000 m, RWT, 16.L1970, wet sclerophyll; 
ANIC (ex. Berl. 999), Wombat Creek, 6 km NE 
Piccadilly Circus, 35°19'S 148°51'E, 750 m, JFL, T. 
Weir and M.-L. Johnson, 30.vi.1984, open forest; 
ANIC (ex. Berl. 1001), Piccadilly Circus, 35°22'S 
148°48'E, 1240 m, JFL, T. Weir and M.-L. Johnson, 
30.vi.1984, subalpine eucalypt litter; ANIC (ex. Berl. 
1000), Blundells Creek, 3 km E Piccadilly Circus, 
35°22'S 148°50'E, 850 m, JFL, T. Weir and M.-L. 
Johnson, 30.vi. 1984, open forest; ANIC (ex. Berl. 821), 
Brindabella Range, Franklin Rd, N end Moonlight 
Hollow, 2 km SW Bulls Head, 35°24'S 148°48'E, M.S. 
Harvey and R.J. Moran, 3.iv.l983; ANIC (ex. Berl. 
926), Ginini Flat, 2 km NE Mt Ginini, 35°31'S 
148°46'E, 1580 m, L. Hill, 20.viii.1983; ANIC (ex. 
Berl. 659), Mt Ginini, 35°32'S 148°46'E, 1660 m, JFL 
and T. Weir, 16.X.1979; ANIC (ex. Berl. 1068), 1 km 
S Mt Ginini, 35°33'S 148°46'E,JFL, ll.xi. 1986; ANIC 
(ex. Berl. 704, 705), 1 km N Mt Gingera, 35°33'S 
148°47'E, A.A. Calder, 18.ii.1981; ANIC (ex. Berl. 
26), Mt Gingera, 35°34'S 148°47'E, c. 5500 ft., E.B. 
Britton, 13.iv.1967, wet sclerophyll; ANIC (ex. Berl. 
50), Mt Gingera, summit, E.B. Britton and Misco, 
19.vii.1967; ANIC (ex. Berl. 661), Mt Gingera, E.C. 
Zimmerman, 20.xi.1979; ANIC (ex. Berl. 830, 831), 
Mt Gingera, 1620-1700 m, L. Hill, 6.iii.l982; ANIC 
(ex. Berl. 1084), Snowy Flat Creek, 0.5 km NE Mt 
Gingera, 35°35'S 148°47'E, A.A. Calder, 28.vi.1988. 

VIC: ANIC (ex. Berl. 1045), Cobb Hill, 14 
km SE Bonang, Goonmirk Ra, 37°18'S 148°50'E, JFL 
and N. Lawrence, 24.xi.1985. 

LORD HOWE ISLAND: AM KS 35592, 
NE area of Mt Gower summit, moss forest near 
campsite, 31°35.2'S 159°04.7'E, 855 m, M.R. Gray, 
12-15.ii.1971; AM KS 35589, creek crossing above 
Boat Harbour, 31°33.5'"S 159°05.5'E, 60 m, M.R. 
Gray, 8.U.1971; AM KS 82998, female (Figs. 6h, 8a, 
d, f, g), KS 82999, male (Figs. 6k, 1, o, 7g, m, 8c, h, 
m), KS 83000, male (Figs. 6m, n, 7c, e, f, i), west end 
of Mt Gower summit on south edge, 31°35.32'S 
159°04.2'E, I. Hutton, 15.V.2001; AM KS 84206- 
84233, additional localities/samples on Mt Gower, AM 
KS 84234-84237, four localities on Mt Lidgbird, I. 
Hutton and CBCR, 2000-2002; AM KS 84238, North 
Hummock, trail to Intermediate Hill, 31°32'54"S 
159°04'58"E, CBCR, 3.xii.2000, mixed rf; AM KS 
84239, western slope of Malabar Ridge, 31°30'57"S 
159°03'31"E, CBCR, 24.xi.2000, broad megaphyllous 
closed sclerophyll forest; AM KS 84240, Transit Hill, 
31°32'01"S 159°04'40"E, I. Hutton, 14.iv.2002; AM 
KS 84241, Little Island, below Far Flats, 31°34'08"S 
159°04'32"E, I. Hutton, 10.viii.2001, under Ficus 



Proc. Linn. Soc. N.S.W., 125, 2004 



195 



A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 



columnaris. 

Description 

Length (anterior margin of head shield to 
telson) up to 6.6 mm; length of head shield up to 0.7 
mm; leg 15 33-40% length of body. Colour: head shield 
and maxillipede pale orange; antenna and most tergites 
orange-yellow, T14 and tergite of intermediate 
segment deeper orange; legs 1-13 pale yellow to pale 
orange, legs 14 and 15 may be deeper orange. 

Head shield (Fig. 5a) smooth, of equal length 
and width, slightly wider than Tl, median notch 
contributing to biconvex anterior margin; longitudinal 
median furrow incised to transverse suture, about one- 
third length of head shield; posterior two-thirds of 



region distal to antennocellar suture desclerotised; setae 
on head shield arranged with bilateral symmetry, four 
larger pairs anterior to antennocellar suture, ten pairs 
behind suture, including four evenly spaced 
submarginal pairs; head shield lacking posterior and 
lateral borders. 

Antenna 27-32% length of body, 2.5-3.3 
times length of head shield, composed of 14 or 
(usually) 15 articles; basal two articles enlarged, most 
articles in distal half moniliform, sclerotised part 
generally of subequal length and width; ultimate article 
about twice length of penultimate. Basal article bearing 
about a dozen sensilla microtrichoidea proximally on 
dorsal side (Fig. 6a). Trichoid sensilla arranged in three 
whorls per article; one or occasionally two curved, 





■J if $A£ 





^^ ^1 




Figure 5. a-e, Dichelobius giribeti n. sp. a, AM KS 82636, female, Monga SF, NSW, head shield, maxillipede 
segment and Tl; b-e, AM KS 82630, male, legs 12-15, Badja SF, NSW. All scales 100 urn. 



196 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE 




Figure 6. Dichelobius giribeti n. sp. Scanning electron micrographs, a-g, Badja SF, NSW; h, k-o, Mt 
Gower, Lord Howe Island; i, j, Monga SF, NSW. a-g, AM KS 82631, female, a, cluster of sensilla 
microtrichoidea on proximal part of antenna, dorsal side, scale 10 |im; b, clypeus, scale 50 urn; c, posterior 
part of clypeus and labrum, scale 50 u.m; d, labral margin, scale 10 Jim; e, antennal articles 10-13, dorsal 
side, scale 30 jim; f, basiconic sensillum at anterior edge of antennal article 12, dorsal side, scale 5 urn; g, 
tip of terminal antennal article, scale 10 |im. h, AM KS 82998, female, dental margin of maxillipede, 
scales 100 um, 30 pm. i, j, AM KS 82638, male, dental margin and ventral view of maxillipede, scales 50 
]im, 100 Jim. k, 1, o, AM KS 82999, male, k, porodont, scale 10 |jjn. 1, dental margin of maxillipede, scale 
50 |im. o, anterior angle of telopodite of first maxilla, scale 10 jjjn. m, n, AM KS 83000, male, telopodite of 
maxillipede and detail of tarsungulum, showing sensilla coeloconica, scales 50 u.m, 5 u.m. 



Proc. Linn. Soc. N.S.W., 125, 2004 



197 



A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 



digitiform sensilla near anterior edge on dorsomedial 
side of a few, variable antennal articles (Fig. 6e); four 
or five articles with a single, short, fusiform sensillum 
at anterior edge on dorsal side (Fig. 6f), most consistent 
on articles 11, 12 and 14; digitiform and fusiform 
sensilla sometimes cooccur on a single article (article 
7 or 9); ultimate article with cluster of 8 or 9 trichoid 
sensilla at apex, one or two curved, digitiform sensilla 
behind apical cluster (Fig. 6g). 

Clypeus with apical cluster of three setae on 
ventral side near lateral margin, single seta medially 
(Fig. 6b); transverse band of four setae in front of 
labrum, outer pair slightly to distinctly smaller than 
inner (Fig. 6c); transverse seta projecting from 
sidepiece; labral margin moderately concave where 
cluster of 7-13 bristles projects; bristles with numerous 
short, spine-like projections along lateral margins and 
on ventral surface along their lengths (Fig. 6d). 
Tomosvary organ large, longitudinally ovate, outer 
edge at lateral margin of cephalic pleurite (Fig. 8k). 

Maxillipede (Figs 6h-n): coxosternal width 
across dental margin 39-44% maximum width; lateral 
margin flexed inward at base of dental projections and 
less convergent than against posterior part; each dental 
margin convex, usually with 5+5, 4+5 or 5+4 teeth, 
sometimes 4+4, 6+5, 5+6 or 6+6; inner tooth smaller 
than others, its apex well posterior to base of outer 
tooth; median notch varying from broadly V-shaped 
(Fig. 6h) to deeply parabolic (Fig. 61); porodont of 
similar length and thickness to largest coxosternal 
setae, its socket at posterolateral edge of outermost 
tooth (Fig. 6k); setae relatively sparsely, fairly evenly 
scattered on coxosternite; tarsal and pretarsal parts of 
tarsungulum of about equal length (Fig. 6m). Dorsal 
and ventral sides of tarsungulum with several sensilla 
coeloconica (Fig. 6n). Bands of pleural collar separated 
by longitudinal median suture (Fig. 6j). 

Mandible: Six curved aciculae (Fig. 7j), all 
with many (up to 1 8) short, blunt denticles along both 
margins (Fig. 7i) on distal half to two-thirds. Four 
paired teeth, dorsal three with accessory denticle field 
delimited by deep groove; dorsalmost tooth and basal 
part of second and third teeth composed of densely 
tuberculate rhomboid and polygonal scales (Fig. 71), 
becoming denticulate near furry pad (Fig. 7m). Fringe 
of branching bristles terminates against dorsalmost 
acicula (Fig. 7f); ventralmost bristles in fringe with 
flattened bases lacking spines, distal two-thirds with 
short spines along both margins and on outer face; 
bristles multifurcating at their distal tips, with three or 
four spines that are longer and thicker than those more 
proximally (Figs 7f, k); more dorsal bristles gradually 
become more uniformly spinose to their broader bases, 
with more numerous distal spines (Fig. 7k), grading 



into wide scales that form a nearly continuous double- 
fringe of hair-like spines, each scale composed of a 
narrow outer fringe and a wider inner fringe, each with 
12-15 spines per scale (Fig. 71); fringe terminates at 
edge of dorsalmost tooth, against a large, smooth scale 
that separates dentate lamina from furry pad (Fig. 7m). 
Furry pad composed of a few scales with distal spines 
and cluster of six or seven mostly simple, elongate 
spines. 

First maxilla: sternite indistinctly delimited 
from coxa (Fig. 7a), short, wide. Coxal projections 
tapering, with rounded apex bearing four or five simple 
setae; one small seta along inner margin near base of 
coxal projection. Telopodite strongly delimited from 
coxal projection; basal article of telopodite with single 
marginal seta anterolaterally or lacking setae; distal 
article with one or two setae near outer margin, anterior 
angle terminating as a long, stout spine; entire inner 
margin fringed with row of six or seven plumose setae 
(Fig. 7b), paired in posterior part of row, with slender 
branchings along more than half of their length (Fig. 
7c); five shorter simple setae inserting near bases of 
plumose setae on ventral side; anterior plumose setae 
fringed on dorsal side by a few elongate spines. 

Second maxilla: anterior margin of coxa 
gently concave; band of four or five small setae across 
anterior part of coxa. Inner edge of tarsus with a row 
of five or six brush-like setae with abundant, slender 
branchings nearly to their bases (Fig. 7d, h). Claw 
composed of up to five digits with concave, scoop- 
like inner surfaces (Fig. 7g); large, curved medial digit 
with furrows or sutures running along its length (Fig. 
7e); outer digits shorter, separated from medial digit 
by a slender, spine-like digit. 

Tergites smooth, all with rounded posterior 
angles, lacking projections; Tl about 85% width of 
widest tergite (TT10 or 12). Posterior margins of TT1, 
3, 5 and 7 transverse (Fig. 4a); TT8, 10 and 12 gently 
concave; TT9, 11, 13 and 14 transverse to weakly 
concave; tergite of intermediate segment transverse or 
gently concave, posterior angle rounded. Two or three 
moderately long setae on lateral margins of long 
tergites, usually with short setae between these; 
posterior margins of tergites fringed with four to twelve 
setae, generally more abundant on more posterior 
segments (maximal number typically on T13); setae 
on inner part of long tergites include transverse band 
of up to six setae across anterior third, two or three 
pairs in two bands behind this. 

Legs 12-15 (Fig. 5b-e) with length ratios 1: 
1.1 : 1.3-1.4 : 1.7. Leg 15 basitarsus 85-115% length 
of distitarsus (Fig. 5e); basitarsus 70-75% length of 
tibia; tibia 2.9-3.4 times longer than maximal width, 
basitarsus 3.4-4 times, distitarsus 5.2-5.7 times. 



198 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE 




Figure 7. Dichelobius giribeti n. sp. Scanning electron micrographs. Scales 10 um except where indicated, 
a, b, d, h, j-1, AM KS 82631, female, Badja SF, NSW; c, e-g, i, m, Mt Gower, Lord Howe Island, a, ventral 
view of first maxillae, scale 50 um; b, distal article of telopodite of first maxilla; d, h, tarsus and claw of 
second maxilla, scales 10 um, 30 um; j, aciculae; k, 1, ventral and dorsal parts of fringe of branching 
bristles on mandible, c, e, f, i, AM KS 83000, male, c, plumose setae on inner margin of telopodite of first 
maxilla; e, claw of second maxilla; f, aciculae and fringe of branching bristles on mandible; i, aciculae. g, 
m, AM KS 82999, male, g, claw of second maxilla, scale 10 um; m, dorsalmost tooth of mandible and 
furry pad. 



Basitarsus 90% length of distitarsus on leg 14 (Fig. 
5d). Coxal projections on leg 15 tapering (in ventral 
view) at about 25-30 degrees; terminal spine with 
distinct (Fig. 8e) or indistinct (Fig. 8i) basal joint, its 



surface with fine longitudinal grooves and ridges like 
those on pretarsal claws. Trochanter of leg 15 with 
small ventrodistal spur (Figs 5e, 8h). Prefemur of legs 
14 and 15 with large ventrodistal spur; leg 15 spur 



Proc. Linn. Soc. N.S.W., 125, 2004 



199 



A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 



with basal width about 25% maximum width of 
prefemur (Fig. 4b). Sharp distal spinose projections 
on tibiae of legs 1-11, absent on legs 12-15. Two 
tarsomeres of leg 13 defined by distinct constriction 
in width and weak articulation without flexure; 
articulation between tarsomeres stronger on leg 14. 
Setae fairly evenly distributed on all podomeres along 
leg, tarsal setae only slightly more slender than those 
on prefemur-tibia; proximo-distal gradient in setal 
thickness enhanced on legs 14 and, especially, 15, with 
distinctly thickened prefemoral setae, including on 
dorsal .side of leg. Anterior and posterior accessory 
claws present on all legs, 25-40% length of main claw 
(Fig. 8a, b); accessory claws with closely-spaced linear 
ridges on their surface except for pitted proximoventral 
part separated by a shallow suture (Fig. 8c). Main claw 
curved, subdivided by sutures; deepest sutures define 
an elongate scute on both lateral sides of claw, proximal 
end of this scute at about distal end of shorter accessory 
claw; large pore or pair of pores at proximal end of 
scute on both sides of leg (Fig. 8c); strong suture 
extends from lateral pore across ventral surface of main 
claw (Fig. 8d), defining proximal end of an elongate, 
triangular ventral scute (Fig. 8g). Proximal part of main 
claw densely pitted; on ventral side of claw, ornament 
changes abruptly at suture delimiting lateral scute, 
becoming linear grooves and ridges as on accessory 
claws (Fig. 8d), with these lineations well developed 
on lateral scute and along length of claw on dorsal 
side; change from pitted to linear ornament gradual 
on dorsal side of claw, with pits irregular proximally, 
becoming aligned as rows of pits, then linear grooves. 
Pair of distally-directed spines proximoventrally, at 
distal end of a curved suture (Fig. 8d); larger spine not 
more than not more than one-eighth length of main 
claw, with tiny subsidiary spine at its base (Fig. 8b). 

Coxal pores: on legs 14 and 15; 2,2/2,2 in 
females (Fig. 4b), 1,1/1,1 in small males, either 1,1/ 
1,1 or 2,2/2,2 (Fig. 4c) in large males, occasionally 
one and two pores on opposing sides of either leg or 
1,2/1,2; pores round, separated by less than their 
diameter when paired; inner pore often smaller than 
outer pore in male, inner pore sometimes larger than 
outer pore in female. 

Female (Fig. 4b): Sternite of segment 15 
gently convex posteromedially, fringed by a 
submarginal setal band that extends along entire 
posterolateral and posterior margin; several setae 
scattered on inner part of sternite. Posterior margin of 
first genital sternite moderately embayed between 
gonopod articulations, sternite bearing 6-11 setae. 
Gonopod with pair of spurs at terminus of a short (Fig. 
8n) to moderately long (Fig. 8e, f) projection; bases of 
spurs nearly touching each other; inner spur 



substantially shorter and narrower than outer spur, both 
bullet-shaped, pointed (Fig. 8n); four or five setae on 
basal article of gonopod, three large setae on second 
article, one large seta on third (Fig. 8j); second and 
third articles variably with one and two smaller setae, 
respectively, on ventromedial face (Fig. 8n); claw 
simple. 

Male (Fig. 4c): Posterior margin of sternite 
15 evenly convex; 10-13 setae fringing margin of 
sternite, 10-12 additional setae scattered over its ventral 
surface; first genital sternite entire medially, bearing 
6-12 setae aligned in two imprecisely-defined 
transverse rows; gonopod bearing two or three setae 
on first article, two on second article, none or one on 
third article, which grades into long, flagelliform 
terminal process, up to 80% length of rest of gonopod 
(Fig. 81); terminal process bearing numerous slender 
spines proximally (Fig. 8m). 

Larvae: five larval stadia (ANIC Berl. 18 and 
231) identified as LO-LIV by comparison to limb 
development in other Lithobiomorpha (Table 1). LI 
with 1 1 antennal articles; LII-LF/ all with 14 articles. 
LII and LIII with 2+2 teeth on dental margin of 
maxillipede; LIV with 3+3 teeth. 

Discussion 

Specimens from Lord Howe Island resemble 
those from the Australian mainland in all meristic 
characters and in fine detail. Intrapopulation variation 
is observed with respect to the number of teeth on the 
maxillipede coxosternal margin, the depth of the 
median notch in the maxillipede coxosternite 
(relatively shallow in Fig. 6h, relatively deep in Fig. 
61), the concavity of the posterior margins of the short 
tergites, and the length of the spur-bearing process on 
the female gonopod. Samples vary in the frequency 
with which large males have either one or two coxal 
pores on legs 14 and 15 (usually two in Lord Howe 
specimens versus one in the large sample from Clyde 
Mountain, NSW, but also two in large specimens from 
the type locality and in the Brindabella Range, e.g., 
Piccadilly Circus, Mt Gingera and Mt Ginini). 

Distinction from other congeners is indicated 
in key above. Dichelobius relictus and D. giribeti are 
consistently distinguished by the presence of a spiracle 
on segment 14 in the former, and D. relictus is 
generally a deeper brown colour. The two species share 
minute details of mandibular and maxillary structure, 
indeed to the extent that description of the mouthparts 
for D. giribeti serves for D. relictus as well. 

The early larval stadia of Dichelobius giribeti 
differ in detail from those of Lithobiidae and 
Henicopinae (see Table 1) with respect to limb 
development. Segmentation of L0 is matched by 



200 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE 




Figure 8. Dichelobius giribeti n. sp. Scanning electron micrographs, a, c, d, f-h, m, Mt Gower, Lord Howe 
Island; b, e, Monga SF, NSW; i-1, n, Badja SF, NSW. a, d, f, g, AM KS 82998, female, a, pretarsus of leg 
14, scale 10 pm. d, g, ventral views of pretarsus of leg 14, scales 10 Jim; f, gonopods, scale 30 Jim. b, e, AM 
KS 82637, female, b, pretarsus of leg 14, posterior view, scale 10 Jim; e, ventrolateral view of first genital 
sternite and gonopods, scale 100 |jm. c, h, m, AM KS 82999, male, c, pretarsus of leg 15, detail of anterior 
accessory claw, scale 5 |im; h, prefemur of leg 15, anterior side, scale 100 |0,m; m, terminal process on 
gonopod, scale 10 |xm. i, j, n, AM KS 82632, female, i, leg 15 coxal process, scale 30 |jm; j, n, lateral and 
ventral views of gonopod, scales 50 jxm, 10 |jm. k, AM KS 82631, female, cephalic pleurite with Tomosvary 
organ, scale 50 Jim. 1, AM KS 82633, male, gonopod, scale 30 Jim. 



Proc. Linn. Soc. N.S.W., 125, 2004 



201 



A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 



Table 1. Comparison of limb development in larval stadia of Lithobiomorpha. Modified from Andersson 
(1979:TabIe II), adding data for Dichelobius giribeti. 





Lamyctes 

emarginatus 
Lithobius 8 spp. 


Lamyctes 
coeculus 


Dichelobius 
giribeti 


Pairs of 


Pairs of 


Pairs of 


Stadium 

LO 
LI 
Lll 
Llll 
LIV 


half- 
Legs developed Limb-buds 
legs 
7 1 large 

7 1 2 small 

8 2 

10 2 
12 _ — 3 


half- 
Legs developed Limb-buds 
legs 

6 2 large 

6 2 

8 2 

10 2 

10 Q 


half- 
Legs developed Limb-buds 
legs 


d £. large 
6 2 2 

8 2 

10 2 

12 3 


\£. o 



Lamyctes coeculus, but larval stadium LI has a unique 
combination of half-developed legs and limb-buds in 
D. giribeti. Segmentation of stadia LII-IV is as in other 
lithobiomorphs. Four larval stadia identified by Eason 
(1993) for Anopsobius macfaydeni have seven, eight, 
ten and twelve pairs of legs, the last three obviously 
being LII-LIV. The taxonomic significance of the 
distinction between six- and seven-legged first larval 
stages in Dichelobius giribeti and Anopsobius 
macfaydeni is unclear without additional data for 
Anopsobiinae. 

Dichelobius bicuspis Ribaut, 1923 

Dichelobius bicuspis Ribaut, 1923:24, Figs. 27-34. 
Dichelobius bicuspis: Wiirmli, 1974:526. 

Material 

NEW CALEDONIA: PROV- NORD: AM 

KS 83001, 1 female, 1 male, Mt Panie, nr summit, 
20°34'S 164°46'E, 1500 m, C. Burwell, 9.xi.2001, rf; 
MNHN, 1 female, 1 larval stadium LIV, Mt Panie, 
20°34'53"S 164°45'38"E, 1350 m, J. Chazeau, A. & 
S. Tillier, 18.xi.1986, wet Agathis forest; QM S60653, 
1 female, Pic d'Amoa, N slopes, 20°58'S 165°17'E, 
500 m, GBM, 10.xi.2001, rf; QM S60654, 1 male, Me 
Maoya, summit plateau, 21°12'S 165°20'E, 1400 m, 
GBM, 1 2.xi.2002, rf. PROV. SUD: MNHN, 3 females, 
Mt Do, 21°45'37"S 165 59'33"E, 840 m, A. & S. 
Tillier & Monniot, 2.iv. 1987, wet Araucaria forest; 
QM S60655, 1 male, Mt Humboldt refuge, 21°53'S 
166°24'E, 1300 m, GBM, 7-8.xi.2002, rf; AM KS 
83002, 1 male, R Bleue, Pourina Track, 22°04'S 
166°38'E, 900 m, GBM, 1 8.xi.2001 , rf; AM KS 83003, 
1 male, Mt Ouin, 22°01'S 166°28'E, 1100 m, GBM, 
9.xi.2002. rf; AM KS 83004, 1 female, 1 male, QM 



S60656, 1 male, Mt Mou base, 22°05'S 166°22'E, 200 
m, GBM, 30.X.2001, 15.xi.2001, rf; MNHN, 3 females, 
1 juvenile, Riviere Bleue, 22°06'13"S 166°39'16"E, 
160 m, A. & S. Tillier, Lviii.1986-30.iv.1987; QM 
S60657, 1 male, Mt Koghis, 22°11'S 166°01'E, 750 
m, GBM, 29.xi.2000, rf; AM KS 83005, 1 female, 
Yahoue, 22°12'S 166°30'E, 100 m, GBM, 4.xi.2001, 
rf. 

Remarks 

Dichelobius bicuspis was based on a few 
specimens from Mt Humboldt (the type locality) and 
Mt Canala, New Caledonia, with Wiirmli (1974) 
adding a record at Nekliai. New collections are listed 
above to indicate that the species has a more 
widespread distribution. 



ACKNOWLEDGEMENTS 

I thank Suzanne Bullock (Scientific Interface) and Sue 
Lindsay and Yongyi Zhen (Australian Museum) for 
assistance with illustrations and electron microscopy, and 
the referees for useful suggestions. Geoff Monteith 
(Queensland Museum) and Jean-Paul Mauries (Museum 
National d'Histoire Naturelle, Paris) kindly provided material 
from New Caledonia. Collection study was hosted and loans 
were arranged by Matthew Colloff (Australian National 
Insect Collection), Gonzalo Giribet and Laura Leibensperger 
(Harvard University), and Mark Harvey and Julianne 
Waldock (Western Australian Museum). Verena Stagl 
(Naturhistorisches Museum Wien) is thanked for arranging 
loan of C. Attems' types. 



202 



Proc. Linn. Soc. N.S.W., 125, 2004 



G.D. EDGECOMBE 



REFERENCES 

Andersson, G. (1979). On the use of larval characters in 
the classification of lithobiomorph centipedes 
(Chilopoda, Lithobiomorpha). In 'Myriapod 
Biology' (Ed M. Camatini) pp. 73-81. 
(Academic Press: London). 

Archey, G. (1917). The Lithobiomorpha of New Zealand. 
Transactions and Proceedings of the New 
Zealand Institute 49, 303-318. 

Archey, G. (1937). Revision of the Chilopoda of New 
Zealand. Part 2. Records of the Auckland 
Institute and Museum 2, 71-100. 

Attems, C. (1911). Myriopoda exkl. Scolopendridae. Die 
Fauna Sudwest-Australiens. Ergebnisse der 
Hamburger sudwest-australischen 
Forschungsreise 1905 3, 147-204. Gustav 
Fischer: Jena. 

Attems, C. (1928). The Myriapoda of South Africa. 

Annals of the South African Museum 26, 1-431. 

Attems, C. (1938). Die von Dr. C. Dawydoff in 
Franzosisch Indochina Gesammelten 
Myriopoden. Memoires du Museum national 
d'Histore naturelle, Paris 6, 2, 187-353. 

Chamberlin, R.V. (1920). The Myriopoda of the 

Australian region. Bulletin of the Museum of 
Comparative Zoology at Harvard College 64, 1- 
269. 

Chamberlin, R.V. (1962). Chilopods secured by the Royal 
Society Expedition to southern Chile in 1958- 
59. University of Utah Biological Series 12, 1- 
23. 

Eason, E.H. (1964). 'Centipedes of the British Isles'. 
(Frederick Warne & Co.: London). 

Eason, E.H. (1993). A new species of Anopsobius from 
the Falkland Islands, with commentary on the 
geographical distribution of the genus 
(Chilopoda: Lithobiomorpha). Myriapodologica 
2, 83-89. 

Edgecombe, G.D. (2001). Revision of Paralamyctes 
(Chilopoda: Lithobiomorpha: Henicopidae), 
with six new species from eastern Australia. 
Records of the Australian Museum 53, 201-241. 

Edgecombe, G.D. (2003). A new species of the 

Gondwanan centipede Anopsobius (Chilopoda: 
Lithobiomorpha) from New South Wales, 
Australia. Zootaxa 204, 1-15. 

Edgecombe, G.D. (2004). The henicopid centipede 

Haasiella (Chilopoda: Lithobiomorpha): new 
species from Australia, with a morphology- 
based phylogeny of Henicopidae. Journal of 
Natural History 38, 37-76. 



Edgecombe, G.D. and Giribet, G. (2003). Relationships of 
Henicopidae (Chilopoda: Lithobiomorpha): new 
molecular data, classification and biogeography. 
African Invertebrates 44, 13-38. 

Edgecombe, G.D., Giribet, G. and Wheeler, W.C. (2002). 
Phylogeny of Henicopidae (Chilopoda: 
Lithobiomorpha): a combined analysis of 
morphology and five molecular loci. Systematic 
Entomology 27, 31-64. 

Mesibov, R. (1986). 'A Guide to Tasmanian Centipedes.' 
(Privately published: Zeehan, Tasmania). 

Ribaut, H. (1923). Chilopodes de la Nouvelle-Caledonie 
et des lies Loyalty. In 'Nova Caledonia. 
Recherches scientifique en Nouvelle-Caledonie 
et aux lies Loyalty. A. Zoology 3(1)' (Eds F. 
Sarasin and J. Roux) pp. 1-79. (C.W. Kreidel's 
Verlag: Berlin, Weisbaden). 

Shinohara, K. (1982). A new genus of centipede of the 
subfamily Anopsobiinae (Henicopidae, 
Chilopoda). Proceedings of the Japanese 
Society of Systematic Zoology 24, 41-46. 

Silvestri, F. (1899). Contribution al estudio de los 
quilopodos chilenos. Revista Chilena de 
Historia Natural 3, 141-152. 

Silvestri, F. (1909a). Contribuzioni alia conoscenza dei 
Chilopodi. in. Descrizione di alcuni generi e 
specie di Henicopidae. Bolletino del 
Laboratorio di Zoologia generate e agraria, 
Portici 4, 38-50. 

Silvestri, F. (1909b). Descrizioni preliminari di vari 

Arthropodi, specialmente d' America. V. Nuovi 
genere di Henicopidae (Chilopoda). Rendiconti 
delta R. Accademia dei Lincei 18, 270-271. 

Silvestri, F. (1933). Nuovi contributi alia conoscenza della 
fauna delle isole italiane dell'Ageo. I. 
Descrizione di un nuovo genere di Chilopodo 
Henicopino. Bolletino del Laboratorio di 
Zoologia generate e agraria, Portici 27, 58-60. 

Verhoeff, K.W. (1925). Systematik. C. Supplement zu den 
Lithobiomorpha. In 'Klassen und Ordnungen 
des Tier-Reichs' (Ed H.G. Bronn) pp. 595-603. 
(Akademische Verlagsgesellschaft: Leipzig). 
Verhoeff, K.W. (1939). Von Dr. G. H. Schwabe in Chile 
gesammelte Isopoda terrestria, Diplopoda und 
Chilopoda. Archiv fur Naturgeschichte, 
Zeitschrift fur Wissenschaftliche Zoologie 
Abteilung B 8, 301-324. 

Wiirmli, M. (1974). Ergebnisse der Osterreichischen 

Neukaledonien-Expedition 1965. Chilopoden. 
Annalen des Naturhistorisches Museums in 
Wien 78, 523-533. 

Zalesskaja, N.T. (1975). New genera and species of 

Chilopoda (Lithobiomorpha) from central Asia 
and Far East. Zoologicheskii zhurnal 54, 1316- 
1325. 



Proc. Linn. Soc. N.S.W., 125, 2004 



203 



204 



A Survey of Ectoparasite Species on Small Mammals During 
Autumn and Winter at Anglesea, Victoria 

Hay lee J Weaver 1 * 2 and John G Aberton 1 . 

'School of Ecology and Environment, Deakin University, Geelong VIC 3217; 2 Present address: School of 
Biological and Environmental Sciences, Central Queensland University, Rockhampton QLD 4702 

(h.weaver@cqu.edu.au) 



Weaver, H.J. and Aberton, J.G. (2004). A survey of ectoparasite species on small mammals during autumn 
and winter at Anglesea, Victoria. Proceedings of the Linnean Society of New South Wales 125, 205-210. 

A survey of the ectoparasites of small native mammals was carried out between April and August 2002, 
in heathlands surrounding Anglesea, Victoria. Antechinus minimus, A. agilis, Rattus lutreolus, R. fuscipes, 
Sminthopsis leucopus and Isoodon ob&sulus were the dominant host mammal species examined. A total of 
921 ectoparasites were collected and identified as five flea species, seven mite species and two species of 
tick. Isoodon obesulus was found to have the highest ectoparasite species richness, with eleven of the 
fourteen species present; while 5. leucopus displayed the lowest ectoparasite species richness with only 
three species found on the hosts examined. The flea Pygiopsylla hoplia was the only ectoparasite species in 
this study to have a distribution across all host mammal species. A new distribution record was made for a 
Haemaphysalis tick species. 

Manuscript received 16 October 2003, accepted for publication 8 January 2004. 

KEYWORDS: Anglesea, ectoparasites, host specificity, marsupials, rodents, species richness. 



INTRODUCTION 

The main groups of ectoparasitic arthropods 
encountered on Australian mammals include fleas 
(order Siphonaptera), mites (order Acariformes), ticks 
(order Parasitiformes) and lice (order Phthiraptera). 
These ectoparasites, as a group, have evolved 
specialised piercing and sucking mouthparts, designed 
for the extraction of blood from a host, with the degree 
of host specificity displayed by ectoparasites varying 
amongst species (Kemp et al. 1982; Dunnet and 
Mardon 1991). 

Many species of ectoparasites are of 
considerable medical and veterinary importance. Fleas 
are capable of transmitting various rickettsial, filarial 
and protozoan diseases (Dunnet and Mardon 1991), 
and ticks can transmit pathogenic filariae, bacteria, 
protozoa, rickettsiae and viruses to wild and domestic 
animals and humans (Obenchain and Galun 1982; 
Aeschlimann 1991). Previous research on ectoparasites 
in the Anglesea region has been limited to flea surveys 
as a precursor to the introduction of myxomatosis 
(Dunnet and Mardon 1991) and calicivirus (F. 
Bartholomaeus pers. comm.), and basic natural history 
of ticks (Roberts 1970). Ectoparasites also negatively 
impact on the health of both domestic and wild animals 
through large infestations, which are of importance in 



management considerations of rare or endangered 
small mammal species present at Anglesea as increases 
in host densities may increase ectoparasite loads. 

The objective of this study was to survey 
ectoparasite species on small native mammals near 
Anglesea, Victoria because an awareness of the 
ectoparasites is important for the potential transmission 
of disease to humans, domestic animals and livestock. 
It is also of interest to the general ecology of small 
mammals in the region. 



METHODS 

Ectoparasites were removed from small 
mammals trapped at two sites at Anglesea, Victoria 
(Fig. 1). The sites chosen for study were the Eumeralla 
Scout camp (38°24'0"S, 144°12'36"E) and Bald Hills 
Road (38°23'24"S, 144°8'24"E) at the Alcoa Lease. 
Both sites were selected using knowledge that they 
contained many host species, and these species were 
all relatively abundant. The Eumeralla Scout camp 
consisted of a coastal tea tree, Leptospermum 
continentale shrub layer, with plants varying from 20 
centimetres to over two metres in height and 
Eucalyptus obliqua at a height of over two metres 



ECTOPARASITES ON SMALL MAMMALS 




Alcoa Lease 
boundary — ►/ 


Bald Hills Rd 1 


1 ! 
i Y Eumera " a f** 

>t L Scout CampC 






yp^ Anglesea 


Great Ocean 


Rd /y 

' Airey's Inlet 


N 

1 

12 3 4km 




scale 



Figure 1. Location map of study area. 



forming the canopy. The site was a flat open heathland 
with woodland dispersed through it, and a swamp 
consisting mainly of Gahnia radula and also L. 
continentale. The Bald Hills Rd site on the Alcoa Lease 
was situated on a slope of approximately 30° in a 
southwesterly direction. The heathland was dominated 
by L. continentale, Epacris impressa, Conospermum 
mitchelli, L. myrsinoides, Platylobium obtusangulum 
and G. radula were the main species present in the 
understorey. Stands of Eu. willisi and Banksia 
marginata were present at the study site. 

Trapping of small mammals was carried out 
during Autumn and Winter 2002, due to the study 
being an honours project requiring completion during 
an academic year. Trapping sessions of three nights 
each were carried out at Eumeralla in June (3-6.6.02), 
July (8-1 1 .7.02) and August (6-9.8.02), with a total of 
1 3 1 mammals captured over the three sessions and at 
the Bald Hills Rd site in April/May (29.4-2.5.02), July 
(22-25.7.02) and August (19-22.8.02), with 161 



captures recorded. Any previously 
trapped mammals captured again in 
following sessions were re-examined 
for ectoparasites and were counted 
accordingly. Fifty aluminium Elliott 
traps (32 x 9 x 10 cm) were placed in 
transects across the Eumeralla site. 
The site at Bald Hills Rd consisted of 
100 traps set in a grid pattern (100 m 
x 100 m) at ten metre intervals. Traps 
were baited using a rolled oats, peanut 
butter and honey mix and were cleared 
within three hours of sunrise. 

Upon capture, mammals 
were transferred from the trap into a 
lightweight mesh bag, identified, ear 
notched for identification purposes, 
weighed, sexed and inspected for 
ectoparasites. As ticks were physically 
attached to the host, they were 
removed using fine forceps to grip the 
tick as close to the host's skin as 
possible and flipping it over to remove 
the tick while leaving the mouthparts 
intact. Fleas and mites were removed 
by ruffling the host's pelage with 
fingers in order to dislodge the 
ectoparasites, or the host was combed 
using Licemeister combs or animal 
flea combs. Numbers of each 
ectoparasite taxa were recorded from 
each mammal and all ectoparasites 
collected were placed in labelled 
containers of 70% ethanol. 

Identification of fleas, mites 

and ticks were carried out using descriptions provided 

by Dunnet and Mardon (1974), Domrow (1987, 1991) 

and Roberts (1970) respectively. 

A linear regression on host mammal body 

weight and ectoparasite species richness was carried 

out using log transformed data. 



RESULTS AND DISCUSSION 

A total of 292 individual mammals were 
trapped over 1350 trap nights from the two sites. The 
host mammals trapped included Antechinus minimus 
(74), A. agilis (69), Sminthopsis leucopus (4) 
(Dasyuriomorphia: Dasyuridae), Isoodon obesulus 
(10) (Peramelemorphia: Peramelidae), Rattus fuscipes 
(50) and R. lutreolus (85) (Rodentia: Muridae). 

Examination of 296 host mammals yielded 
364 fleas and 557 acari (mites and ticks) in total. From 
this, five flea species were identified, along with seven 



206 



Proc. Linn. Soc. N.S.W., 125, 2004 



H.J. WEAVER AND J.G. ABERTON 



mite species and two tick species. Of these, two species 
of mites were unable to be identified to species level; 
these were referred to by their family names as 
unidentified Laelapidae and unidentified 
Trombiculidae. Table 1 shows the number of 
examinations of each host mammal species and the 
species of ectoparasites removed from the host species. 

The most common host examined for 
ectoparasites was Rattus lutreolus, with 85 
examinations and the host examined least was 
Sminthopsis leucopus with only four examinations. 
Sminthopsis leucopus is an uncommon mammal in the 
Anglesea area. Lunney (1995) states although it has a 
wide distribution throughout southern Australia, it 
prefers sparse ground to forage, whereas the sites in 
this study had very dense ground cover. 

Figure 2 shows /. obesulus as having the 
greatest ectoparasite species richness and S. leucopus 
the smallest. A significant linear association was found 
between host weight and ectoparasite species richness 
(MS=0.098, F=7.966, df=l, P=0.048) with 64.32% of 
the variation in ectoparasite species richness accounted 
for by mean body weight of the hosts. This is consistent 
with previous studies showing that host body size 
determines ectoparasite species richness (Kuris et al. 
1980, cited in Stanko et al. 2002). Another factor that 
can influence ectoparasite species richness is the social 
behaviour of the host. Stanko et al. (2002) found that 
higher host densities generally equated to lower species 
richness on individuals, possibly because of anti- 
parasitic behaviours such as grooming. As bandicoots 
have a reputation of 'pugnacious behaviour between 
conspecifics' (Lobert 1990) and indicate a low social 
tolerance (Thomas 1990), it could be that the 
bandicoots examined in this study had a higher species 
richness of ectoparasites and a higher abundance of 
each species in part due to a combination of larger 
body size and lack of social grooming. 

The most common ectoparasite collected was 
the flea Pygiopsylla hoplia, which was recorded on 
every host mammal species. According to Dunnet and 
Mardon (1974), P. hoplia is the most commonly 
collected Australian species of flea. It has a distribution 
across Australia, excluding the Northern Territory, and 
has been recorded on many species of peramelids, 
dasyurids and rodents (Dunnet and Mardon 1974). In 
contrast, Stephanocircus dasyuri was mostly recorded 
on /. obesulus, and occasionally on A. minimus. The 
similar foraging nature of both these mammal species 
may be the reason why this species of flea was not 
recorded on any other hosts. Macropsylla hercules was 
only recorded on Rattus spp. and /. obesulus, perhaps 
due to the size of the host animals, as this flea is very 
large. Macropsylla hercules is commonly collected 



from various native Rattus species from southern 
Australia (Dunnet and Mardon 1974). The other 
species of flea collected, Acanthopsylla rothschildi 
rothschildi and Bibikovana rainbowi appeared to 
display little host specificity, as they were recorded 
from the majority of the host species. 

Host specificity for acarine ectoparasites 
collected varied. The highly host specific Androlaelaps 
marsupialis was only found between the groove of the 
tibia and fibula on the hind legs of /. obesulus where 
grooming is difficult (pers. obs.). Similarly, 
Mesolaelaps anomalus was recorded only on /. 
obesulus. In contrast, the trombiculid mites and the 
tick Ixodes tasmani showed a broad host range, being 
found on all host species except for S. leucopus and A. 
minimus respectively. The trombiculids were found 
most frequently inside the ears of hosts during this 
study, but can be found on any exposed skin including 
legs, feet and tails (pers. obs.). Trombiculid rnites are 
parasitic during their larval stage and later live in the 
soil as free living adults (Domrow 1962). One small 
infestation was recorded in the pouch of a female /. 
obesulus, and it has been suggested that larval 
trombiculids occurring in the pouches of A. minimus 
can directly infest any pouch young present (B. Wilson, 
Deakin University, pers. coram.). Ixodes tasmani is a 
common species of tick with a distribution widespread 
across southern Australia with a wide range of hosts 
(Roberts 1970). 

The species of Haemaphysalis collected from 
/. obesulus was identified as H. humerosa, but 
differences in the spiracular plate between the Anglesea 
specimens and specimens from known populations in 
Queensland have been observed. An alternative 
identification is H. ratti. Further research is being 
carried to provide a definite identification of the 
specimens (I. Beveridge, University of Melbourne, 
pers. comm., D. Kemp, CSIRO, pers. comm). 

Other ectoparasitic arthropods were collected 
from host mammals studied. Lice (Phthiraptera, species 
unknown) were collected from R. lutreolus on three 
occasions; but were not observed on any other host 
mammals examined. The rove beetle species 
Myotyphlus jansoni (Coleoptera: Staphylinidae) was 
collected from Rattus lutreolus on five occasions. 
However, M. jansoni is not an obligate ectoparasite. 
Myotyphlus jansoni has only been recorded on a very 
small number of individual native Rattus species 
previously (Hamilton-Smith and Adams 1966). The 
beetles are usually collected near the anus or tail (as 
they were in this study) and have also been recorded 
in bat guano in a cave near Warrnambool, Victoria; 
thus it may be assumed that the beetles feed on the 
excreta of the rats, which is not strictly an ectoparasitic 



Proc. Linn. Soc. N.S.W., 125, 2004 



207 



ECTOPARASITES ON SMALL MAMMALS 






05 



o 

P 

as 



rs 
©, 

S" 
rs 

re 

a 

o 

3 

cr 
o 



' s 

ca 

O 


a 


3 


r 


c 


a 


B". 


a" 

3 


B- 


§- 


ff 


§ 

(T 


3 

a 
si- 


8" 
o 


1 

Co 


3 


ft 


(T> 


S3' 


o 


Co 


O. 


*~-« 


3 


s 


§* 


ft 


<T 


Co 


H 


►2 


1 




a 


■"a 

Co 


O 








o 








o 









o 
o 

1+ 

-J 

oo 



&a 



'3 & 

ft ^s. 



s a- 

co ft 
B 



to H> t> 

S: ft ^ 

S* a o§ 

a*! I" 



I- 



i^ 



a- 



■§ 



a- 



M H (O (O « 

-&>■ o 

o 



5s s? tf S: 5; ^ 

«i a a to to w. 

3 ft* S a 1 5 

- Co S* S" C" a 

S a a ■ g- 




H M Ifl W Ul H 
I— H Ul O 

^3 



to 
to 

1+ 

OO 



a ot> 
s S' 
a- "a 

O .'o 
I - 

1 a" 

a" g* 

3- a" 

% 
a- 



fr 



a s 



a 3 
St 
s 

Co 

**^ 

& 


Rattusfu 
(bush rat 


Antechin 
(agile an 




ft 




o 


"a - 


sT a 


& 


ft 

Co 


B' <*> 


Co 




£ sr* 



oo 
oi 






lit 

a 3 * J 

r §. a 

2 a "-a 

k 2 a 

ft* "5 



o\ o oo 



M w H w w 
O tO © 



On 

o 



SO 


t— ' 


U\ 


o 


1+ 


On 


to 


1+ 


>— * 


to 




oo 



a 
ft 

I 

4a 



S3 
s* o§ 

S* a" 

2 '-a 

1 3 



a- 



a~ 5" 

| a' 



•o on *^ 




OS 



H- 

VO 



ft V 

a oq 

I 5' 

a- ^3 

a .*» 
I - 
^ a" 

a" g" 

pS 

a- a 



u> i— > 



Co 



a oo u ^ 



1? £ 

I I 

p ft 

5 <"> 

I i 

§■§• 
s- a. 

§ I 



-J 


3 

&3 


58 

p 

2, 




rs 


3 




& 


55* 


o 


9 


H 

o 



1+ 



•8 si 

a s 



a S 

2. g 

* S 

0, g- 

a. <*• 

S a 

3 



S5 ^ 
a oo 

a- "« 
a- a 

Co 

ft 

a- 



U m U u 



I s 


a 
a, 


1 
ft 

Co 


H 

3 

1 


a 
a. 


1 

Co 


a 
a. 


ft' 


S 
5" 


O 

a* 
ft 


3 


a 

5" 

ft 


a 
ft 


R- £ 


a* 

Co 




ft 


1" 

Co 


a 
S 

Co 


a 


Co 




p 


Co 






<^ 


ft 


1 


C8 


s. 


1 


s 




ft 


a' 

»** 
a- 




ft" 


a' 
3- 


1 




ft 

a- 


1 

to 




ft 
a- 


1 

Co 


1% 

2" 




K 






K 








Co 


to' 




to 


Co' 





ffl 



s ^ 

1+ ffi 

9? 



2 1 
■o" 

cr 

© 

B 
P 
T3 

J? 

1 
P 



208 



Proc. Linn. Soc. N.S.W., 125, 2004 



H.J. WEAVER AND J.G. ABERTON 



Oh 

3 

• i-H 

a 
a, 

e2 



14 - 

12 

10 

8 

6 

4 

2 






SI 



y = 0.017x + 5.722 
R 2 = 0.643 







i r i i i i i i 

50 100 150 200 250 300 350 400 



Mean body weight (g) 

Figure 2. Relationship between ectoparasite species richness and body weight of host mammals. SI = 
Sminthopsis leucopus, Aa = Antechinus agilis, Am = Antechinus minimus, Rl = Rattus lutreolus, Rf = Rattus 
fuscipes, Io = Isoodon obesulus. 



relationship (Hamilton-Smith and Adams 1966; 
Lawrence and Britton 1991). 

The ectoparasite species collected during this 
study were all considered to be common throughout 
the region (Roberts 1970; Dunnet and Mardon 1974) 
and all are theoretically able to transmit pathogens to 
animals or humans. Generally, fleas are known to be 
intermediate hosts for the cosmopolitan rodent 
tapeworm, Hymenolepis diminuta and the canine 
tapeworm Dipylidium canium, along with being able 
to transmit various filarial, rickettsial and protozoan 
pathogens, however native Australian fleas have not 
been found to contribute epizootics in the field (Dunnet 
and Mardon 1991). Ixodes tasmani has been recorded 
as an intermediate host of various rickettsiae, including 
Rickettsia australis, the organism which causes 
Queensland tick typhus (Campbell and Domrow 1974, 
cited in Cavanagh 1999). Haemaphysalis ticks are 
vectors of Coxiella burnetii (Q fever), in bandicoots 
and macropods and domestic livestock (Kettle 1995). 
Therefore it is recommended that care be taken when 
in areas where ticks are present, especially at the 



Eumeralla Scout Camp where groups of scouts may 
come into contact with ticks while carrying out 
activities in the area. 

In conclusion, it was found that there was a 
significant relationship between ectoparasite species 
richness and body weight of host mammal species. 
There was no difference in the species of ectoparasites 
collected from both study sites, except for M. jansoni, 
which was only found on R. lutreolus at the Bald Hills 
Rd site. As there have been no other studies carried 
out of this type in the region, it is recommended that a 
study over a longer time frame be carried out in order 
to accurately assess seasonal variations of ectoparasite 
numbers. 



ACKNOWLEDGEMENTS 

The authors are grateful to F. Bartholomaeus 
(South Australian Research and Development Institute) and 
M. Shaw (University of Queensland) for additional flea and 
mite identification respectively, and Dr Ian Beveridge 
(University of Melbourne) and Dr David Kemp (CSIRO) 



Proc. Linn. Soc. N.S.W., 125, 2004 



209 



ECTOPARASITES ON SMALL MAMMALS 



for assistance with the identification of Haempahysalis sp. 
We appreciate Philip Barton's comments on the manuscript. 
The study formed part of the principal author's BEnvSc 
Honours thesis and was carried out in accordance with 
Deakin University Animal Ethics committee guidelines and 
conducted under Victorian Department of Natural Resources 
and Environment wildlife permit no. 10001759. 



REFERENCES 

Aeschlimann, A. (1991). Ticks and Disease: Susceptible 
■ hosts, reservoir hosts and vectors. In 'Parasite-host 
associations, coexistence or conflict?' (Eds C.A. 
Toft, A. Aeschlimann and L. Bolis) pp. 148-156. 
(Oxford University Press: Oxford). 

Campbell, R.W. and Domrow, R. (1974) Rickettsioses in 
Australia: Isolation of Rickettsia tsutsugamushi 
and R. australis from naturally infected hosts. 
Transactions of the Royal Society of Tropical 
Medicine and Hygiene 68, 397-402. 

Cavanagh, F.A. (1999) The common marsupial tick, 
Ixodes tasmani, and factors that influence the 
degree of infestation of the common brushtail 
possum, Trichosurus vulpecula. BSc Honours 
thesis, University of Sydney, Sydney. 

Domrow, R. (1962). The mammals of Innisfail. 2: Their 
mite parasites. Australian Journal of Zoology 2, 
268-307. 

Domrow, R. (1987). Acari Mesostigmata parasitic on 
Australian vertebrates: An annotated checklist, 
keys and bibliography. Invertebrate Taxonomy 1, 
817-948. 

Domrow, R. (1991). Acari Prostigmata parasitic on 

Australian vertebrates: An annotated checklist, 
keys and bibliography. Invertebrate Taxonomy 4, 
1283-1376. 

Dunnet, G.M. and Mardon, D.K. (1974). A monograph of 
Australian fleas (Siphonaptera). Australian 
Journal of Zoology Supplementary Series 30, 1- 
273. 

Dunnet, G.M. and Mardon, D.K. (1991). Siphonaptera. In 
'The insects of Australia: a textbook for students 
and research workers Vol II.' (CSIRO) pp. 705- 
716. (Melbourne University Press: Melbourne). 

Hamilton-Smith, E. and Adams, D.J.H. (1966). The 
alleged obligate ectoparasitism of Myotyphus 
jansoni (Mathews) (Coleoptera: Staphylinidae) 
Journal of the Entomological Society of 
Queensland 5, 44-45. 

Kemp, D.H., Stone, B.F. and Binnington, K.C. (1982). 
Tick Attachment and Feeding: Role of the 
Mouthparts, Feeding Apparatus, Salivary Gland 
Secretions and the Host Response. In 'Physiology 
of Ticks'. (Eds F.D. Obenchain and R. Galun) 
pp.1 19-168. (Pergamon Press: Oxford). 

Kettle, D.S. (1995) 'Medical and Veterinary Entomology 
(2 nd edn.)'. (CAB International: United Kingdom). 

Kuris, A.M., Blaustein, A.R. and Alio, J.J. (1980). Hosts 
as islands. American Naturalist 116, 570-586. 



Lawrence, J.F. and Britton, E.B. (1991). Coleoptera. In 
'The insects of Australia: a textbook for students 
and research workers Vol II.' (CSIRO) pp. 543- 
683. (Melbourne University Press: Melbourne). 

Lobert, B. (1990). Home range and activity period of the 
Southern brown Bandicoot {Isoodon obesulus) in 
a Victorian heathland. In 'Bandicoots and Bilbies.' 
(Eds J.H. Seebeck, P.R. Brown, R.L. Wallis and 
CM. Kemper) pp. 319-325. (Surrey Beatty and 
Sons: Sydney). 

Lunney, D. (1995). White-footed Dunnart. In 'The 

mammals of Australia.' (Ed. R. Strahan) pp.143- 
145. (Australian Museum/Reed New Holland: 
Sydney). 

Obenchain, F.D. and Galun, R. (1982). Preface. In 

'Physiology of ticks.' (Eds F.D. Obenchain and R. 
Galun) pp. vii-ix. (Pergamon Press: Oxford). 

Roberts, F.H.S. (1970). 'Australian ticks.' (CSIRO: 
Melbourne). 

Stanko, M., Miklisova, D., Goiiy de Bellocq, J. and 

Morand, S. (2002). Mammal density and patterns 
of ectoparasite species richness and abundance. 
Oecologica 131, 289-295. 

Thomas, L.N. (1990). Stress and population regulation in 
Isoodon obesulus (Shaw and Nodder). In 
'Bandicoots and bilbies.' (Eds J.H. Seebeck, P.R. 
Brown, R.L. Wallis and CM. Kemper) pp. 335- 
343. (Surrey Beatty and Sons: Sydney). 



210 



Proc. Linn. Soc. N.S.W., 125, 2004 



Occurrence and Conservation of the Dugong (Sirenia: 
Dugongidae) in New South Wales 

Simon Allen 1 , Helene Marsh 23 and Amanda Hodgson 23 

Graduate School of the Environment, Macquarie University, NSW 2109; 2 School of Tropical Environment 
Studies and Geography, James Cook University, Townsville, Qld 4811; 3 CRC Reef Research Centre, PO Box 

772, Townsville, Qld 4810 



Allen, S., Marsh, H. and Hodgson, A. (2004). Occurrence and conservation of the dugong (Sirenia: 
Dugongidae) in New South Wales. Proceedings of the Linnean Society of New South Wales 125, 211- 
216. 

Recent sightings of dugongs well beyond the southern limit of their accepted range (~27°S) on the Australian 
east coast prompted a review of past records of dugongs and their current conservation status in New South 
Wales. While archaeological analyses have identified bones of Dugong dugon in Aboriginal middens at 
Botany Bay (~34°S) and colonial records indicate stranded animals as far south as Tathra (~36.5°S), there 
were no verified sightings of live individuals in NSW waters for some years; however, five separate sightings 
of individuals and pairs were documented in the austral summer of 2002/03 in estuaries on the NSW central 
coast (~32-33.5°S). It is suggested that conditions such as warm sea temperatures and low rainfall (promoting 
seagrass growth) may be facilitating explorative ranging south by dugongs. 

The IUCN lists dugongs as 'vulnerable' at a global scale and they are also classified 'vulnerable' under 
the Threatened Species Conservation Act NSW 1995, yet they are not routinely considered in risk assessments 
for inshore development in this State. Threatening processes such as shark meshing persist. The importance 
of considering dugongs in future impact assessments for inshore marine and estuarine developments is 
emphasized. 

Manuscript received 17 October 2003, accepted for publication 8 January 2004. 

KEYWORDS: conservation, distribution, dugong, Dugong dugon, risk assessment, sightings, status, 
vulnerable. 



INTRODUCTION 

The dugong (Dugong dugon), along with all 
other extant Sirenians, is regarded as a shallow water, 
tropical and sub-tropical species (Martin and Reeves 
2002; Rice 1998). Dugongs are thought to be strictly 
marine, inhabiting the coasts of some 37 countries and 
territories (Marsh et al. 2002). Despite their widespread 
distribution, dugong numbers have declined in most 
of their known range and they are believed to be 
represented by fragmented, relic populations in most 
countries. Likely causes for this decline and continuing 
threats include: large-scale destruction of seagrass as 
a result of sedimentation, dredging, mining, trawling, 
and pollution; incidental take as by-catch in 
commercial and recreational gill and mesh nets as well 
as shark nets set for bather protection; direct takes from 
indigenous hunting, and vessel strikes and disturbance 
(Marsh et al. 1999, 2002; Hodgson 2003). 

Australian waters are the dugong' s 
stronghold, where their distribution is described as 
extending from Shark Bay in Western Australia (25 °S) 
around northern Australia to Moreton Bay in southern 
Queensland (27°S) (Marsh et al. 2002). Dugongs are 



a 'listed marine species' under the Australian 
Environment Protection and Biodiversity Conservation 
Act 1999 (EPBC Act). The EPBC Act reflects 
Australia's commitments under various international 
conventions including the Bonn Convention on the 
Conservation of Migratory Species of Wild Animals, 
which lists the dugong on Appendix 2. Dugongs are 
also considered 'vulnerable' under the Threatened 
Species Conservation Act NSW 1995 and under the 
Nature Conservation Act Qld 1992. 

Evidence of a decline in dugong numbers 
along the urban coast of Queensland (Marsh et al. 
2001) led to the establishment of a series of dugong 
protection areas in some key dugong habitats in 
Queensland (Marsh et al. 1999; Marsh 2000). No 
similar protection has been afforded dugongs in NSW, 
presumably on the assumption that only vagrants of 
the species range into NSW waters. Dugongs have been 
considered in some impact assessments for aquaculture 
developments in NSW (e.g. Anon. 2001a), but not 
others (e.g. Anon. 2001b). These assessments occurred 
in the same location, suggesting consideration of 
dugongs and potential impacts thereon is inconsistent 
in NSW. 



DUGONGS IN NEW SOUTH WALES 



In this paper, we highlight past and present 
evidence that the dugong's range on the east coast of 
Australia extends into NSW waters, including 
estuaries, when environmental conditions are suitable. 
Given their conservation status under both international 
conventions and national acts, we suggest that 
occasional visitation warrants adherence to the legal 
obligation of considering dugongs and their preferred 
habitats in future impact assessments. 



EARLY RECORDS TO RECENT SIGHTINGS 

Dugong bones have been found associated 
with edge-ground hatchet heads in Aboriginal middens 
near Sydney, indicating that at least small numbers of 
dugongs have utilized NSW waters for many centuries 
(Etheridge et al. 1896). In 1799 Flinders described the 
catching of dugongs by Aborigines in Moreton Bay, 
southeast Queensland (Mackaness 1979). Aborigines 
in NSW also caught dugongs in more recent times, 



with bones having been found in middens as far south 
as Botany Bay in the late 18 th Century (Troughton 
1928). 

There are currently two sources of dugong 
sightings in NSW: the Atlas of NSW Wildlife and 
records of by-catch from shark meshing supervised 
by NSW Fisheries. The Atlas of NSW Wildlife yields 
83 reports of live, stranded and dead animals for the 
period 1788 to 2003 (Anon. 2003b; Fig. 1). 

A significant portion of these reports (63) 
occurred in late 1992 and throughout 1993. This influx 
of animals occurred after the loss of 1 ,000 km 2 of 
seagrass from Hervey Bay in southeast Queensland 
following floods (Preen and Marsh 1995). Two 
dugongs were caught in NSW shark meshing during 
this time (Swansea in November 1992 and January 
1993). Three earlier captures were also made in shark 
nets (Bronte in July 1951, Bondi in July 1951, 
Queenscliff in April 1971) (Krogh and Reid 1996). 

Only two records of dead and stranded 
individuals have been reported to the NSW National 




Figure 1. Past records of dugongs on the NSW coast from 1788 to 2003 (open circles; Anon. 2003b) and 
dugong sightings in central NSW estuaries during summer 2002/03 (filled circles). 



212 



Proc. Linn. Soc. N.S.W., 125, 2004 



S. ALLEN, H. MARSH AND A. HODGSON 



Estuary 


Date 


Lat./Long. 


Description 


Source 


Wallis 
Lake 


Late Oct. 
2002 


32°11.0' 
152°30.2' 


Kayak tour operator reports dugong/s over 
seagrass beds within Wallis Lake 


S.Smith, 
pers. comm. 


Port 
Stephens 


10 th Jan. 
2003 ' 


32°42.8' 
152°06.7' 


Dolphin watch operators report two adult 
dugongs near Manton Bank 


D. Aldritch, 
pers. comm. 


Lake 
Macquarie 


24 th Jan. 
2003 


33°20.5' 
150°29.8' 


Recreational fishers report cow-calf pair 
travelling seaward out Swansea Channel 


B. Roche, 
pers. comm. 


Port 
Stephens 


1 st Feb. 
2003 


32°41.8' 
152°03.2' 


Dolphin watch operator report dugong/s in 
upper estuary west of Soldiers Point 


D. Aldritch, 
pers. comm. 


Brisbane 
Water 


3 td Feb. 
2003 


33°30.1' 
152°20.3' 


Resident reports dugong/s off Orange Grove 
beach 


Anon. 2003b 



Table 1. Dugong sightings in central NSW estuaries in the austral summer of 2002/2003. 



Parks and Wildlife Service (NPWS) in the last decade, 
with no live sightings occurring until late 2002/03. 
Between late October 2002 and early February 2003, 
five separate sightings of individuals and pairs within 
(or swimming out of) central coastal estuaries were 
reported to NPWS and/or the authors (Table 1; Fig. 
1). These occurred along a c. 200km stretch of coastline 
and we do not know if these sightings include repeat 
sightings of the same individual(s). 

SEAGRASS DISTRIBUTION AND WATER 
TEMPERATURES 

All the estuaries in which dugongs were 
sighted are known to support seagrass meadows (Table 
2). Dugongs have been recorded eating the seagrasses 
listed in Table 2, with the exception of Ruppia spp. 
(Anderson 1986, Marsh et al. 1982, Lanyon et al. 
1989). Species of the genus Halophila are preferred. 
The distribution of dugongs has been reported as being 
constrained to water temperatures >~18°C (Anderson 
1986, 1994; Marsh et al. 1994; Preen et al. 1997). 
However, the water temperatures at the sites in Table 
2 were above this thermal threshold in summer 2002/ 
03. 

DISCUSSION 

The low abundance of dugongs in NSW 
waters may be the result of a number of factors 



including limited availability of seagrass in the region, 
relatively low water temperatures during winter months 
and in open coastal waters between estuary and bay 
habitats, and/or human pressures. The entire NSW 
coast supports only 155 km 2 of seagrass (West et al. 
1989), the major portion of which would be Posidonia 
australis and species of the Zosteraceae family, which 
are not favoured by dugongs. In relative terms, the 
amount of seagrass in NSW is much less than the total 
area of seagrass in Moreton Bay alone (250 km 2 : Abal 
et al. 1998) and would contain correspondingly small 
cover of Halophila spp. Troughton (1928) interpreted 
historical records as suggesting that dugongs may have 
occurred in greater numbers in NSW prior to European 
settlement. It has also been suggested (MacMillan 
1955) that dugong populations on the tropical east coast 
were again beginning to expand into the northern rivers 
region of NSW. Any expansion of the dugong' s range 
into NSW waters further south than this region may 
have been inhibited by the loss of seagrass beds in 
areas such as Port Macquarie and Botany Bay to 
anthropogenic influences (Pointer and Peterkin 1996). 
The dugong observations in 2002/03 (Table 
1) were in areas of NSW which have some of the largest 
seagrass beds, at least two of which include Halophila 
species - part of the preferred diet of dugongs (Marsh 
et al. 1982; Table 2). The increasing evidence that 
individual dugongs embark on movements over many 
hundreds of kilometres within tropical waters (N. Gales 
pers. comm; Marsh and Lawler 2001, 2002; Marsh 



Proc. Linn. Soc. N.S.W., 125, 2004 



213 



DUGONGS IN NEW SOUTH WALES 



Estuary (latitude) 


Seagrass species and approximate area coverage 


Water temp. (<€) 


Wallis Lake 
(~32.2°S) 


Zosteraceae, Posidon ia australis , Ruppia and Halophila 
spp. -30. 785km 2 


October mean: 18.9 
October 2002: 21.0 


Port Stephens 
(~32.7<5) 


Zosteraceae, Posidonia australis and 
Halophila spp. -7 .453km 2 


January mean: 24.1 
February mean: 24.6 


Lake Macquarie 
(-SSI'S) 


Zosteraceae, Posidonia australis, Ruppia and Halophila 
spp. -13. 391km 2 


January mean: 21.6 


Brisbane Water 
(~33.4<5) 


Zosteraceae, Posidonia australis and 
Halophila spp. -5.490km 2 


February mean: 22.1 



Table 2. Extent of seagrass meadows and water temperatures at sighting ocations. Sources for seagrass 
coverage and water temperature data: West et al. (1985) and Anon. (2003a) respectively. Water 
temperatures are means from 1987-2002, unless otherwise stated. 



and Rathbun 1990; Marsh et al. 2002) suggests it is 
possible that dugongs explore and utilize these southern 
seagrass beds. Warm water temperatures during the 
summer months of 2002/03 may have encouraged this 
behaviour. 

Although only five dugongs have been 
reported drowned in shark nets in NSW over the last 
c. 50 years (Krogh and Reid 1996), such deaths are 
not inconsequential since few dugongs are commonly 
found south of Moreton Bay. Two of these mortalities 
coincided with a seagrass dieback event (Preen and 
Marsh 1995) and further impact on Queensland 
seagrass beds or increase in water temperature in NSW 
may see an increase in shark net capture of dugongs 
off NSW beaches. Such events will highlight negative 
effects on populations of non-target species, and the 
efficacy of shark control programs for bather protection 
in NSW and Queensland will again be called into 
question (Anon. 2002). 

The dugong is classified as 'vulnerable' at a 
global scale on the IUCN Red List of Threatened 
Species. As the only extant species in the family 
Dugongidae, the extinction of the dugong will result 
in biodiversity loss at the family and generic levels as 
well as at the species level. In the light of 
inconsistencies evident in risk assessments for inshore 
development in NSW, we re-iterate that dugongs 
should be considered occasional visitors to NSW 
coastal waters. Their limited numbers warrant the 
dugongs' consideration in future impact assessments 
for estuarine and inshore marine developments. The 
estuarine nature of recent sightings suggests that 



explorative ranging by dugongs is not necessarily 
limited to strictly marine environments, rather to areas 
where seagrass beds occur. This also adds weight to 
the importance of assessing potential impacts on 
seagrass habitats. 



ACKNOWLEDGMENTS 

We gratefully acknowledge all those that provided 
prompt and unambiguous reports of recent sightings. We 
would also like to thank Mick Murphy of Hunter Coast Area 
NSW National Parks and Wildlife Service for providing 
access to the relevant wildlife database and Jeanine Almany 
for information on seagrass distribution and water 
temperatures in NSW. SA was supported by an ARC SPIRT 
grant, HM and AH by funding from the CRC Reef Research 
Centre. This manuscript was greatly improved by comments 
from Robert Williams, John Merrick and two anonymous 
reviewers. 



REFERENCES 

Abal, E.G., Dennison, W.C. and O'Donohue, M.H. 

(1998). Seagrasses and mangroves in Moreton 
Bay. In 'Moreton Bay and Catchment' (Eds I.R. 
Tibbetts, N.J. Hall and W.C. Dennison) pp. 269- 
278. (University of Queensland: Brisbane). 

Anderson, P. K. (1986). Dugongs of Shark Bay, Australia 
- seasonal migration, water temperature, and 
forage. National Geographic Research 2, 473- 
490. 



214 



Proc. Linn. Soc. N.S.W., 125, 2004 



S. ALLEN, H. MARSH AND A. HODGSON 



Anderson, P. K. (1994). Dugong distribution, the seagrass 
Halophila spinulosa, and thermal environment 
in winter in deeper waters of eastern Shark Bay, 
Western Australia. Wildlife Research 21, 381- 
388. 

Anon. (2001a). 'Environmental Impact Statement for a 
Commercial Snapper Farm Proposed for 
Providence Bay, NSW. Wildthing 
Environmental Consultants, Raymond Terrace, 
NSW. 

Anon. (2001b). 'Port Stephens Pearl Oyster Industry 
Environmental Impact Statement, Volumes 1 
and 2'. Report No. 1412/R01/V2. Umwelt 
(Australia) Pty Ltd, Toronto. 

Anon. (2002). Nomination of 'Death or injury to marine 
species following capture in beach meshing 
(nets) and drumlines used in Shark Control 
Programs' as a Key Threatening Process under 
the Environment Protection and Biodiversity 
Conservation Act 1999. pp. 20. August. 
Environment Australia, Canberra. Available at 
http://www.ea.gov.au/biodiversity/threatened/ 
nominations/public-comment/shark- 
meshing.html (accessed Sept. 2003). 

Anon. (2003a). Australian Oceanographic Data Centre. 
Available at http://www.aodc.goc.au/ (accessed 
Sept. 2003). 

Anon. (2003b). 'The Atlas of New South Wales Wildlife 
- Fauna'. Accessed April 2003. 

Etheridge, R.J., David, T.W.E. and Grimshaw, J.W. 
(1896). On the occurrence of a submerged 
forest, with remains of the dugong, at Sheas 
Creek, near Sydney. Journal and Proceedings of 
the Royal Society of New South Wales 30, 158- 
185. 

Hodgson, A.J. (2003). The risk of boat disturbance and 
boat strikes to dugongs: observations using 
blimp-cam. In 'Australian Marine Science 
Association Annual Conference' (Eds J.N.A. 
Hooper, N. Hall and B.M. Degnan). July 9 th - 
11 th , 2003, Brisbane. 

Krogh, M. and Reid, D. (1996). Bycatch in the protective 
shark meshing programme off south-eastern 
New South Wales. Biological Conservation 77, 
219-226. 

Lanyon, J., Limpus, C.J. and Marsh, H. (1989). Dugongs 
and turtles: Grazers in Seagrass systems. In 
'Biology of Seagrasses: A treatise on the 
biology of seagrasses with special reference to 
the Australia Region' (Eds A.W.D. Larkum, 
A.J. McComb and S.A. Shepard) pp. 610-634. 
Elsevier, Amsterdam. 

Mackaness, G. (1979). 'The discovery and exploration of 
Moreton Bay and the Brisbane River' . (Review 
Publications: Dubbo). 

MacMillan, L. (1955). The Dugong. Walkabout 

(Australian National Travel Association) 21, 17- 
20. 

Marsh, H. (2000). Evaluating management initiatives 

aimed at reducing the mortality of dugongs in 
gill and mesh nets in the Great Barrier Reef 



World Heritage Area. Marine Mammal Science 
16, 684-694. 

Marsh, H. and Lawler, I. (2001). 'Dugong Distribution 
and Abundance in the Southern Great Barrier 
Reef Marine Park and Hervey Bay: Results of 
an Aerial Survey in October-December 1999'. 
Great Barrier Reef Marine Park Authority 
Research Publication 70. Great Barrier Reef 
Marine Park Authority, Townsville. 

Marsh, H. and Lawler, I. R. (2002). 'Dugong Distribution 
and Abundance in the Northern Great Barrier 
Reef Marine Park - November 2000'. Great 
Barrier Reef Marine Park Authority Research 
Publication 77. Great Barrier Reef Marine Park 
Authorty, Townsville. 

Marsh, H. and Rathbun, G.B. (1990). Development and 
application of conventional and satellite radio- 
tracking techniques for studying dugong 
movements and habitat usage. Australian 
Wildlife Research 17, 83-100. 

Marsh, H, Channells, P. W., Heinsohn, G. E. and 
Morissey, J. (1982). Analysis of stomach 
contents of dugongs from Queensland. 
Australian Wildlife Research 9, 55-67. 

Marsh, H, De'ath, G., Gribble, N. and Lane, B. (2001). 
'Shark Control Records Hindcast Serious 
Decline in Dugong Numbers off the Urban 
Coast of Queensland'. Great Barrier Reef 
Marine Park Authority Research Publication No. 
70. Great Barrier Reef Marine Park Authority, 
Townsville. 

Marsh, H, Eros, C, Corkeron, P. and Breen, B. (1999). A 
conservation strategy for dugongs: implications 
of Australian research. Marine and Freshwater 
Research 50, 979-990. 

Marsh, H, Penrose, H., Eros, C. and Hugues, J. (2002). 
'Dugong Status Report and Action Plans for 
Countries and Territories'. Early Warning and 
Assessment Report Series. UNEP/DEWA/ 
RS.02-1. 162 pp. 

Marsh, H, Prince, R.I.T., Saalfeld, W.K. and Shepherd, R. 
(1994). The distribution and abundance of the 
dugong in Shark Bay, Western Australia. Wildlife 
Research 21, 149-61. 

Martin, A.R. and Reeves, R.R. (2002). Diversity and 

Zoogeography. In Marine Mammal Biology: an 
evolutionary perspective (Ed. A.R. Hoelzel). 
Blackwell Science Ltd. Oxford. 

Pointer, I. R., and Peterken, C. (1996). Seagrasses. In 'The 
State of the Marine Environment Report for 
Australia' (Eds L. P. Zann and P. Kailola.). 
Technical Annex: 1. Great Barrier Reef Marine 
Park Authority, Townsville. 

Preen, A. and Marsh, H. (1995). Response of dugongs to 
large-scale loss of seagrass from Hervey Bay, 
Queensland, Australia. Wildlife Research 22, 
507-519 

Preen, A.R., Marsh, H, Lawler, I.R., Prince, R.I.T. and 
Shepherd, R. (1997). Distribution and 
abundance of dugong, turtles, dolphins and 
other megafauna in Shark Bay, Ningaloo Reef 



Proc. Linn. Soc. N.S.W., 125, 2004 



215 



DUGONGS IN NEW SOUTH WALES 



and Exmouth Gulf, Western Australia. Wildlife 
Research 24, 185-205. 

Rice, D.W. (1998). 'Marine Mammals of the World: 
Systematics and distribution'. Special 
Publication No. 4. The Society for Marine 
Mammalogy. 

Troughton, E.L. (1928) Sea-Cows: The Story of the 

Dugong. The Australian Museum Magazine 3, 
220-228. 

West, R.J., Larkum, A.W.D. and King, R.J. (1989). 

Regional studies - seagrasses of south eastern 
Australia. In 'Biology of Seagrasses: A treatise 
on the biology of seagrasses with special 
reference to the Australian region' (Eds A.W.D 
Larkum, A.J. McComb and S.A. Shepherd). 
Elsevier, Amsterdam. 

West, R.J., Thorogood, C.A., Walford, T.R. and Williams, 
R.J. (1985). An estuarine inventory of New 
South Wales, Australia. Fisheries Bulletin 2. 
Department of Agriculture, NSW. 



216 Proc. Linn. Soc. N.S.W., 125, 2004 



Captures, Capture Mortality, Age and Sex Ratios of Platypuses, 

Ornithorhynchus anatinus, During Studies Over 30 Years in the 

Upper Shoalhaven River in New South Wales 

T.R. Grant 

School of Biological, Earth and Environmental Sciences, University of NSW, NSW 2052 

Email t.grant@unsw.edu.au 



Grant, T.R. (2004). Captures, capture mortality, age and sex ratios of platypuses, Ornithorhynchus 
anatinus, during studies over 30 years in the upper Shoalhaven River in New South Wales. 
Proceedings of the Linnean Society of New South Wales 125, 217-226. 

Data collected during a number of studies over a period of 30 years in the upper Shoalhaven River, New 
South Wales, are presented. A total of 700 individual platypuses (Ornithorhynchus anatinus) were captured 
during the studies, consisting of 137 juvenile females, 94 juvenile males, 292 adult females and 177 adult 
males. The overall sex ratios of both adults (1.65:1) and juveniles (1.46:1) were significantly biased towards 
females. Females were found to live up to 21 years. Very few recaptures of juvenile males made estimates 
of longevity equivocal, but three individuals were at least 7 years old when last captured. Capture and 
handling mortality during the various studies was low (0.86%). Sixty-two percent of platypuses marked in 
the study area were never recaptured, fewer adult males were recaptured than females (36% and 51% 
respectively) and recaptures of juveniles were much lower than for adults (32% females and 14% males). 
Recapture data suggest considerable mobility by adults and dispersal by juvenile platypuses along the upper 
Shoalhaven River and its tributaries. 

Manuscript received 2 September 2003, accepted for publication 8 January 2004. 

Keywords: Age, Capture, Marking, Mortality, Ornithorhynchus anatinus, Platypus , Sex Ratio 



INTRODUCTION 

Over the past 30 years a number of research 
projects has been carried out in a study area in the 
upper Shoalhaven River in New South Wales. 
Individual projects have included investigation of 
temperature physiology (Grant and Dawson 1978a,b; 
Grant 1983; Hulbert and Grant 1983a,b), diet (Faragher 
et al. 1979), movements and home ranges (Grant 1983, 
1992), haematology and pathology (Munday et al. 
1998; Whittington and Grant 1983, 1984, 1995; 
Whittington et al. 2002), lactation and milk 
composition (Grant et al. 1983, Gibson et al. 1988; 
Grant and Griffiths, 1992) and population genetics 
(Gemmell 1994; Akiyama 1999). During these studies, 
long-term data have been collected on recaptures, 
capture mortality, longevity and sex ratios of 
platypuses (Ornithorhynchus anatinus). 

During the later studies from 1987, the 
investigation and use of Passive Integrated 
Transponder (PIT) tags or "microchips" was begun, 
probably the first time that this marking method was 
used on a wild mammalian species in Australia (Grant 
and Whittington 1991). The long-term success of this 



method in the mark and recapture studies of the 
platypus is reported below. 

Collins (1973) tabulated the ages of eight 
platypuses kept in captivity in a variety of locations, 
including the Bronx and Budapest Zoos. These ranged 
from four to 17 years, although anecdotal information 
from zoos and sanctuaries indicates that the species 
may survive in captivity for up to 21 years (Whittington 
1991). Concerning the longevity of platypuses in the 
wild, the naturalist Harry Burrell (1927) wrote that "the 
length of life of the platypus is not known. It is my 
intention to ring-mark some fully furred young as 
opportunity offers, and it may be that we shall gain 
some information on this point at a later date, if these 
marked individuals are captured". Burrell did not later 
report the ages of platypuses he may have "ring- 
marked". However, Grant and Griffiths (1992) 
reported the ages of platypuses marked in the upper 
Shoalhaven River as being between as much as 4 years 
for males and 8 for females. Since that report, a further 
12 years of research has resulted in the data presented 
in this paper on the ages and sex ratios in this 
population of platypuses. Mortality of capture and 
handling of platypuses in previous studies has not 



CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN PJVER 



previously been discussed in the literature and this 
aspect of the studies is presented in the current paper. 



METHODS 

Between June 1973 and January 2004, 
platypuses were captured in 16 pools in the upper 
Shoalhaven River in New South Wales using the 
unweighted "gill" net methods outlined in Grant and 
Carrick (1974). Until 1987, individuals were marked 
using stainless steel leg bands (Grant and Carrick 
(1974) but these were phased out after trials on the 
use of Passive Integrated Transponder (PIT) tags 
proved to be successful (Life Chip tags; Destron 
Fearing Corporation Scanner; Grant and Whittington 
1991). 

Sex was determined using the presence or 
absence of the adult spur or the morphology of juvenile 
spurs. Absolute ages were determined from individuals 
initially captured as juveniles and minimum ages for 
adults were estimated using the time of loss of the 
female spur, the morphological changes in the males 
spur (Temple-Smith 1973; Grant 1995), and 
subsequent recaptures. Females possessing a spur were 
categorised as being in their first year of life (0 years 
of age) and males could be assigned to their first or 
second year of life (0 or 1 year of age). As the females 
in this area lose the spur between October and 
December in their first year after emergence from the 
nesting burrows, any female lacking a spur at first 
capture was considered to be = 1 year of age (i.e. in 
their second year of life). It should be noted that two 
female juvenile platypuses bred in captivity at Taronga 
Zoo in the 2002/2003 breeding season apparently lost 
their spurs within only 4 months after emergence from 
the nesting burrow (Adam Battaglia, Taronga Zoo, 
pers. comm.). Males with adult spur morphology were 
considered to be at least in their third year of life, or = 
2 years old. Subsequent recaptures permitted minimum 
ages to be assigned to individuals, beginning with a 
minimum age at first capture of one year for adult 



females and two years for adult males (Temple-Smith 
1973; Grant and Griffiths 1992; Grant 1995). 

Recaptures were recorded for animals in all 
of the 16 pools of the 12.5 km section of the upper 
Shoalhaven River and 3.9 km of an adjacent creek. 
However, by 1987 a number of these pools had filled 
with sand and were no longer netted. By 1993, the 
previously largest and deepest pool (1 km long x 2-5 
m deep) was completely filled with sand and was no 
longer sampled, although many of the platypuses 
originally captured in this pool were captured in the 
pools downstream. From 1988, when PIT tagging had 
become the predominant method of marking, sampling 
was mainly restricted to three pools in the Shoalhaven 
River itself and one in the adjacent creek. In most years 
after that time these pools were sampled late in the 
year (mainly December) when lactating animals were 
most likely to be caught and at the end of summer 
(mainly February or March) when juveniles had newly 
emerged (Grant and Griffiths 1992). 



RESULTS 

Sex ratios 

During the studies from June 1973 to January 
2004, 700 individual platypuses were captured. Table 
1 shows the numbers in each of four age/sex classes. 
All sex ratios were significantly biased towards 
females. The ratio of females to males was 1.58 
females: 1 male (Chi 2 = 34.87; p < 0.001) for all 
animals, 1.46:1 for juveniles (Chi 2 = 8.00; p < 0.01) 
and 1.65:1 for adults (Chi 2 = 27.38; p < 0.001). 

Age 

Only 45 individuals (41 females and 5 males), 
first marked as juveniles, were subsequently 
recaptured. Figure 1 shows the distribution of ages of 
these individuals at their latest recapture. Two juvenile 
females were recaptured regularly over periods of 13 
and 16 years but were not captured again in 5-6 
subsequent years. These animals were assumed to have 





Juvenile 


Adult 


Total 


Males 


94 


177 


271 


Females 


137 


292 


429 


Total 


231 


469 


700 


Sex Ratio (F:M) 


1.46:1 


1.65:1 


1.58:1 


Chi 2 


8.00 


27.38 


34.87 


P 


** < 0.005 


** < 0.001 


** < 0.001 



Table 1. Numbers and sex ratios of platypuses captured in the upper Shoalhaven River study area. 
** significant at < 0.01 level. 



218 



Proc. Linn. Soc. N.S.W., 125, 2004 







T.R. 


GRANT 






Year 


Juvenile 


Juvenile 


Year 


Adult 


Adult 


(Actual) 


Female 


Male 


(Minimum) 


Female 


Male 





96 


89 





- 


- 


1 


21 


4 


1 


181 


28 


2 


3 


1 


2 


33 


117 


3 


3 


- 


3 


18 


11 


4 


3 




4 


18 


9 


5 


2 


- 


5 


10 


6 


>5 


9 


- 


>5 


32 


6 


Total 


137 


94 


Total 


292 


177 



Table 2. Numbers of platypuses allocated to actual and estimated minimum age categories in the upper 
Shoalhaven River study area. Actual = ages of animals initially captured as newly-emerged juveniles; 
Minimum = minimum ages; calculated from years between initial and last capture of individuals first 
captured as adults. 



died. However, one was again recaptured and lactating 
at the end of the study, when her age was 21 years. As 
indicated in the Methods section, females without spurs 
are at least in their second year of life (>/=l year old) 
and it is possible to attribute males to either their first 
second or third year of life (0, 1 or 2 years old) based 
on spur morphology changes. Ages of >/= 2 years 
could be attributed to 1 1 1 female adults caught and 
subsequently recaptured. Similarly 32 adult males were 
attributed to the >/= 3 year age category. The 
distribution of these minimum ages are shown in Table 
2 and Fig. 1. 

While most platypuses caught in the study 
could only be attributed to the >/= 1-2 year age 
category, 9 females first captured as juveniles survived 



between 5-21 years and 32 of those initially captured 
as adults survived 5-15 years. One juvenile male was 
subsequently recaptured at 2 years of age but 32 males, 
first captured as adults, survived to minimum ages of 
3-7 years. 

Recaptures 

The numbers of juvenile, adult male and adult 
female platypuses recaptured at least once in the latter 
12 years (when most animals were marked with PIT 
tags) were not significantly different from those of the 
first 18 years of the study (when the majority were 
marked with leg-bands) (Table 3). Table 4 presents 
combined recapture data for both leg-banded and PIT 
tagged animals for the whole study period. 





Juvenile 


Juvenile 


Adult 


Adult 




Female 


Male 


Female 


Male 


Leg-banded 










Total recaptures 


33 


9 


105 


53 


(£ 1 recapture) 










Total captures 


97 


69 


214 


134 


% recapture 


34.0% 


13.0% 


49.1% 


39.5% 


PIT tagged 










Total recaptures 


24 


2 


47 


15 


(> 1 recapture) 










Total captures 


54 


32 


86 


41 


% recapture 


42.5% 


6.3% 


54.7% 


36.6 


Chi z 


1.60 


1.04 


0.77 


0.12 


P 


NS<0.20 


NS <0.31 


NS<0.38 


NS <0.73 



Table 3. Comparison between total number of recaptures (>/= 1) for leg-banded and PIT tagged platypuses 
in the upper Shoalhaven River study area. Animals marked with both bands and PIT tags between 1987 
and 1991 are included in both sets of data. NS = not significant. 



Proc. Linn. Soc. N.S.W., 125, 2004 



219 



CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER 



Juvenile Females 



200 -i 

180 - 

160 - 

> 140- 

g 120- 

01 

3 100 - 

8" 80 - 

* 60- 

40 - 

20 - 





T 



T 



123456789 



— -n 1 1 1 1 1 1 r-~i 

10 11 12 13 14 15 16 17 18 19 20 21 



Age (years) 



Adult Females 




1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 

Age (years) 

Figure 1. Actual and minimum age frequencies of platypuses in the study. Actual ages are shown for 
those animals initially caught as juveniles (Juvenile Females and Juvenile Males). Estimated minimum 
ages shown are for animals caught first as adults (Adult Females and Adult Males). 
CONTINUED ON FACING PAGE. 



Considerable numbers of both male and female adults 
and juveniles were captured only once. 

Table 4 shows that total recaptures (>/= 1 
times) and recaptures in the categories of 1, 2-5 and 
>5 times were lower for adult males than for females 
and that total recaptures of juvenile males was less 



than half (14%) that of juvenile females (32%). The 
majority of recaptured males were caught within the 
first months after initially emerging from the nesting 
burrows (0 years of age), while recaptures of juvenile 
females were spread across 0-21 years after emergence 
(Figure 1 and Table 4). 



220 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT 



Juvenile males 



180 - 
160 
140 - 
& 120 
| 100 

& 60 
40 
20 - 




80 



1 23 4 5 6 7 8 9 10 11 12 13 14 15 16 



Age (years) 



Adult Males 





180 -j 




160 - 




140- 




120 - 


g 

§■ 

to 


100 - 


80 - 
60 - 




40 - 




20 - 




o -1 




1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 

Age (years) 



Capture mortality 

Of the 700 platypuses captured during the 
various research projects six died as a result of capture 
and handling (0.86%). Two drowned as a result of 
netting, two died suddenly within a few hours of 
capture (sudden death; animals appeared healthy and 
no obvious cause of death was identified from post- 
mortem examination by veterinarians), one succumbed 
to anaesthesia and one became caught in submerged 
vegetation by its transmitter attachment and drowned 
during telemetry work. Details are in Table 5. 



DISCUSSION 

Sex Ratios 

As reported by Grant and Griffiths (1992) for 
the first 18 years of the study, the sex ratios for both 
adults and juveniles were significantly biased towards 
females. Table 6 compares the sex ratios for captured 
adult platypuses in three other areas (Grant 
unpublished). Although based on much smaller sample 



Proc. Linn. Soc. N.S.W., 125, 2004 



221 



CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER 





Juvenile 
Female 


Juvenile 

Male 


Adult 
Female 


Adult 
Male 


1 recapture 
2-5 recaptures 
>5 recaptures 
Total recaptures 
(> 1 recaptures) 


17 

19 

7 

43 


10 

3 


13 


66 

58 
18 
142 


34 

20 

6 

60 


% recapture 


32% 


14% 


51% 


36% 


Total captures* 


135 


94 


278 


165 



Table 4. Total recaptures of male and female juvenile and adults platypuses in the upper Shoalhaven 
River study area. Mortalities and some animals which would have been unlikely to have been 
recaptured after the netting of some pools was discontinued are not included in this total captures 
figure. 



sizes, none of these were significantly different from 
parity. Grant and Griffiths (1992) also reported no 
significant difference between males and females in 
total numbers of platypuses (juvenile and adults not 
specified) captured in various rivers of New South 
Wales and the Australian Capital Territory (Table 6). 
Like the situation in the upper Shoalhaven River, 
during the earlier years of a study (1986-90) in the 
Duckmaloi River on the central tablelands of New 
South Wales, a bias towards females in both adult and 
juvenile platypuses was found. However, in the later 
years (1991-2000) more adult males than females were 
recorded (David Goldney, University of Sydney, 
Orange, pers. comm.). 

The recapture data discussed below seems to 
indicate that female platypuses in the upper Shoalhaven 
River survived for significantly longer periods than 
males. Over time, this longer survival of females would 
presumably have led to a sex ratio weighted towards 
females in the population. However, this explanation 
does not account for the disparity between numbers 
of male and female juveniles in this population. 

Most juvenile male platypuses disappeared 



from the upper Shoalhaven River population in their 
first year (86%; Table 4). However, 13% of juvenile 
females were recaptured in the area up to age one year, 
19% were recaptured later than two years after 
emerging from the nesting burrows and two even 
remained in the area up to age 13 and 21 years 
respectively. Twelve juvenile females (9%) bred in the 
area, eight of these over a number of breeding seasons. 
Differential dispersal may contribute to the difference 
in the adult sex ratio, but again this does not explain 
the significant bias to females in the numbers of 
juveniles captured at the time they were becoming 
independent (late January-late March), unless most 
male juveniles dispersed immediately after 
independence, with females dispersing later. 
Unfortunately the data from this study do not permit 
this hypothesis to be rigorously tested, as most 
sampling only occurred early and late each year. 

The possibility also exists that the uneven sex 
ratios are determined by differential fertilisation of 
eggs, development of embryos or pre-emergence 
survival of young but no explanation arises from the 
data collected in this study concerning the significant 



Cause of Death 


Juvenile 
Female 


Juvenile Male 


Adult Female 


Adult 
Male 


Total 


Drowned in net 
Sudden death 
Anaesthesia (ether) 
Snagged transmitter 


1 

1 









1 


1 




2 




2 
2 
1 
1 


Total 


2 





2 


2 


6 



Table 5. Capture and handling mortalities in platypuses during work on various projects in the upper 
Shoalhaven River study area 



222 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT 



Location 


Adult Female 


Adult 
Male 


Sex Ratio 
(F:M) 


Chi z 
Probability 


Various streams NSW/ACr f 
Various streams NSW/ACT* 
Barnard River, NSW 
Thredbo River, NSW 
Wingecarribee River, NSW 


101 

47 
24 
14 
29 


117 
47 
22 
10 
30 


1.15:1 

1:1 
1.10:1 
1.4:1 
1:1.03 


1.17 
p< 0.28 

0.09 
p<0.77 

0.67 
p<0.41 

0.02 
p<0.90 


Shoalhaven River, NSW 


285 


173 


1.85:1 


2738** 
p< 0.001 



Table 6. Comparison of sex ratios of adult platypuses in various studies in New South Wales and the 
Australian Capital Territory (ACT). Collected by Temple-Smith and * Griffiths (from Grant and Griffiths 
1992); ** significant at < 0.01 level. 



bias towards females in the sex ratio of juvenile 
animals. In the Barnard River study referred to in Table 
6, where the adult sex ratio was not different from 
parity, the sex ratio for juveniles was heavily male- 
biased (14 males; 3 females) during the single breeding 
season studied. A similar result was also obtained once 
during a single breeding season (12 males to 3 females; 
1978/79) in the current study, indicating differences 
between individual years. However, in the 24 years 
during the study in which juveniles were captured, the 
numbers of females exceeded males in 87.5% of those 
years. Considerable annual differences in recorded 
annual sex ratios were also found in the Duckmaloi 
River study (David Goldney, University of Sydney, 
Orange pers. comm.). 

Age 

The minimum estimated age for male 
platypuses in this study (7 years) was considerably less 
than for females (up to 21 years), with nine adult 
females surviving for a minimum of 10 years. These 
data suggest that females live longer than males in the 
wild. However, determination of actual age, or the 
estimated minimum age, depended on recapture data 
and, as discussed, recapture of males was much lower 
(36% recaptured >/= once) than for females (51% 
recaptured >/= once). After being regularly captured 
previously, 13 and 21 year old females (first marked 
as juveniles) had not been recaptured for the last 6 and 
5 years respectively of the study. Except for one adult 
female, which had been captured in December 2002 



and was not caught in March 2003, all the other adult 
females >/= 10 years of age had also not been 
recaptured in the latter years of the study. These data 
appeared to indicate a life span of 10-16 years may 
represent an expected upper range of longevity for 
female platypuses in the wild. However, the final 
recapture of one female at the age of 21 years showed 
a maximum female longevity in the wild comparable 
to that in captivity. While the data for males in the 
wild appeared to show shorter life spans (up to 7 years), 
this could equally represent non-recapture of older 
males. Reports do not suggest differing longevity 
between the sexes in captivity (Collins 1973; 
Whittington 1991). 

Recaptures 

PIT tags were initially used because of the 
occurrence of notched and broken male spurs as a result 
of bands abrading the spur base. However, it was also 
suspected that the lower capture rates in males, 
particularly juveniles may have been attributable to 
band losses. Bands were normally fitted more loosely 
to males to permit the much greater radial growth of 
the hind legs in this sex. No significant differences 
between captures for banded and PIT tagged males 
(Table 3) indicated that band loss could not fully 
explain the lack of recaptures, although four female 
animals, initially marked with bands and PIT tags, were 
found to have lost their bands during the latter part of 
the study, indicating some band loss. Only one PIT 



Proc. Linn. Soc. N.S.W., 125, 2004 



223 



CAPTURE, MORTALITY AND SEX OF PLATYPUSES FN THE SHOALHAVEN PJVER 



Home 
range(km) 



Maximum distance 
(km) 



Source 



Location 



0.2-2.0 


5.6 (24 hr max. 


= 4.0) 


Grant, 1983 


Shoalhaven River 


0.4-2.3 


2.3 




Grant, 1983, Grant et al. 1992 


Thredbo River 


0.3-23 


2.3 




Serena, 1994 


Badger Creek 


2.9-7.0 


15.0 




Gardner and Serena, 1995 


Watts River and Badger Creek 


0.4-2.6 


2.6 




Gust and Handasyde, 1995 


Goulburn River 



Adult: 


24 hr max. = 10.4 


Serena et al. 1998 


Yarra River, Mullum Mullum Creek, 


2.9-7.3 


(male) 




Diamond Creek 


Juvenile: 


24hrmax. = 4.0 






1.4-1.7 


(female) 








40 in 18 months 


Australian Platypus 


Yarra catchment 




(juvenile) 


Conservancy 1999 


Andersons to Steels Ck 




48 in 7 months 


Australian Platypus 


Wimmera River 




(young male) 


Conservancy 2001 





Table 7. Home ranges and maximum distances moved by platypuses in various studies, including the 
Shoalhaven River (bold). 



tag failure or loss was confirmed in 220 tags applied 
to animals during the studies. The lack of spur damage, 
some evidence of band loss and no significant 
differences being found between recaptures of leg- 
banded and PIT tagged animals confirmed PIT tagging 
as the preferred method of marking platypuses (Grant 
and Whittington 1991). 

Large numbers of both adult male (64%) and 
female (49%) platypuses were not recaptured in the 
study area after being marked either with leg bands, 
PIT tags or both. This observations suggests one, or a 
combination of the following: 

Loss of marks . Double marking indicated that some 
band loss did occur during the study but there was 
little indication that PIT tags were lost or failed. 

Mortality . Little is known about the causes and 
incidence of mortality in platypuses. 

Foxes (or dogs) will take platypuses on land, 
from shallow riffle areas and by digging into burrows 
(Serena 1994; Grant 1993; Anon. 2002). Large eagles 
may also be possible natural predators of platypuses 
(Rakick et al. 2001). The remains of a platypus near a 
burrow excavated by a fox or dog, an isolated skull in 
a pool and part of a skull in a pile of other mammalian 
bones (mainly cattle and sheep) were the only observed 
evidence of mortality found during the studies in the 
upper Shoalhaven River. 

While 50% of 131 individuals tested positive 
to leptospirosis antibodies {Leptospira interogans 
serovar hardjo)(Mux\ddiy et al. 1998), no clinical 
symptoms of the disease were observed and nothing 
is known of any disease organism, resulting in 
significant mortality in this population. The Mucor 
fungus, which has caused mortality in Tasmanian 



populations, has not so far been detected in mainland 
populations of platypuses, including those in the upper 
Shoalhaven River (Whittington et al. 2002). 

Mobility . Diurnal and longer-term mobility over 
distances of up to 5.6 kilometres have been previously 
reported in individuals in the upper Shoalhaven River 
(Grant 1983, 1992, 1995; unpublished) and in other 
studies. These data are summarised in Table 7. 

After the marking of 700 individual 
platypuses (including significant numbers of new 
juveniles) during the 30 years, it was expected that the 
majority of the population would eventually be marked 
and that unmarked dispersing juveniles or adults might 
still enter the area but would be in fairly small numbers. 
In fact, considerable numbers of new adult animals 
were captured throughout the study. Some individuals 
were captured as many as 20 times over periods of up 
to 21 years and yet the times between recaptures of 
these individuals was often quite variable. For example, 
despite the pools being regularly netted during the 
study, two adult females were only subsequently 
recaptured nine and 10 years respectively after their 
initial capture in those pools. Even for females 
identified as breeding in particular pools during 
different breeding seasons, periods of time between 
some recaptures of these animals ranged from 1 to 10 
years. 

These latter observations suggest that a great 
deal of mobility probably characterises the platypus 
populations in the upper Shoalhaven River, although 
the effects of mortality and/or dispersal cannot be ruled 
out as reasons for the influx of new animals and the 
lack of recapture of a significant proportion of the 
platypuses in the upper Shoalhaven River study area. 
All of these possibilities need further study. 



224 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT 



While there was capture and handling 
mortality during the studies in the upper Shoalhaven 
River population, this was quite low (< 1%) due to the 
utilisation of methods developed through considerable 
experience by the author and other researchers over 
the past three decades. 

ACKNOWLEDGMENTS 

The many friends and colleagues who assisted with 
the field work over the years are too numerous to thank 
individually but all are gratefully acknowledged. The last 
10 years of the study would not have been possible without 
the help and support of Colin, Kate, Sue and Tom Heath 
and Paul Anink, Marie-Louise Lissone and Gina Grant. The 
late Athol MacDonald and the Izzard and Laurie families 
are acknowledged for their permission for access to the river 
and creek, and for their friendship and assistance in various 
aspects of the field work. Unfortunately Bill and Ron Izzard 
both died in 2003 and this paper is dedicated to their memory. 
Adam Battaglia and David Goldney are acknowledged for 
their personal communications. Richard Whittington and 
Joanne Connolly carried out the post-mortem examinations 
of the two animals which died suddenly after capture. Peter 
Temple-Smith, Michael Augee and an anonymous referee 
provided valuable comments on the manuscript. Some of 
the work reported was done while in receipt of funding from 
the Environment Australia (then Australian National Parks 
and Wildlife Service) and the Australian Research Council 
(then Australian Research Grants Committee). This work 
was carried out under NSW National Parks and Wildlife 
Service Scientific Investigations Licence A 184, New South 
Wales Fisheries Scientific Research Permit F84/1245 and 
University of New South Wales Animal Care and Ethics 
Approvals 94/91, 97/46 and 00/45. 



REFERENCES 

Akiyama, S. (1999). Molecular ecology of the platypus 
{Ornithorhynchus anatinus). PhD thesis, La 
Trobe University, Melbourne. 

Anonymous. (1999). Keeping tabs on a marathon 
swimmer. Ripples 113, 1 

Anonymous. (2001). Platypus on the move. Ripples 18, 1 

Anonymous. (2002). Foxes kill four platypus. Hastings 
Gazette. 19 December. 

Burrell, H. (1927). The platypus. Sydney: Angus and 
Robertson. 

Collins, L.R. (1973). 'Monotremes and marsupials. A 
reference for zoological institutions'. 
Smithsonian Institution Press, Washington DC. 

Faragher, R.A., Grant, T.R. and Carrick, F.N. (1979). 
Food of the platypus {Ornithorhynchus 
anatinus) with notes on the food of the brown 
trout (Salmo trutta) in the Shoalhaven River, 
New South Wales. Australian Journal of 
Ecology 4, 171-179. 



Gardner, J.L. and Serena, M. (1995). Spatial organisation 
and movement patterns of adult male platypus, 
Ornithorhynchus anatinus (Monotremata: 
Ornithorhynchidae). Australian Journal of 
Zoology 43, 91-103. 

Gibson, R.A., Neumann, M., Grant, T.R. and Griffiths, M. 
(1988). Fatty acids of the milk and food of the 
platypus {Ornithorhynchus anatinus). Lipids 23: 
377-379. 

Gemmell, N.J. (1994). Population and evolutionary 

investigations in the platypus {Ornithorhynchus 
anatinus): a molecular approach. PhD thesis, La 
Trobe University, Melbourne. 

Grant, T.R. (1983). Body temperature of free-ranging 
platypuses, Ornithorhynchus anatinus, with 
observations on their use of burrows. Australian 
Journal of Zoology 31, 117-122. 

Grant, T.R. (1992). Captures, movements and dispersal of 
platypuses, Ornithorhynchus anatinus, in the 
Shoalhaven River, New South Wales, with 
evaluation of capture and marking techniques. 
In. Platypus and Echidnas. (Ed M.L. Augee). 
pp. 255-262. (Royal Zoological Society of 
NSW: Sydney). 

Grant, T.R. (1993). 'The Bellinger River Water Supply 

Project Aquatic Studies - The Platypus'. Report 
to Mitchell McCotter on behalf of the Coffs 
Harbour City Council and Department of Public 
Works by Mount King Ecological Surveys, 
May, 1993. 

Grant, T.R. (1995). 'The platypus. A unique mammal'. 
2nd edition. Sydney: University of NSW Press: 
Sydney 

Grant, T.R. and Carrick, F.N. (1974). Capture and 
marking of the platypus, Ornithorhynchus 
anatinus, in the wild. Australian Zoologist 18: 
133-135 

Grant, T.R. and Dawson, T.J. (1978a). Temperature 
regulation in the platypus, Ornithorhynchus 
anatinus, maintenance of body temperature in 
air and water. Physiological Zoology 51:1-6. 

Grant, T.R. and Dawson, T.J. (1978b). Temperature 
regulation in the platypus, Ornithorhynchus 
anatinus, production and loss of metabolic heat 
in air and water. Physiological Zoology 51: 315- 
332. 

Grant, T.R. and Griffiths, M. (1992). Aspects of lactation 
and determination of sex ratios and longevity in 
a free-ranging population of platypuses, 
Ornithorhynchus anatinus, in the Shoalhaven 
River, New South Wales. In. Platypus and 
Echidnas. (Ed M.L.Augee). pp. 80-89. (Royal 
Zoological Society of NSW: Sydney). 

Grant, T.R., Griffiths, M. and Leckie, R.M.C. (1983). 
Aspects of lactation in the platypus, 
Ornithorhynchus anatinus (Monotremata), in 
waters of eastern New South Wales. Australian 
Journal of Zoology 31: 881-889. 



Proc. Linn. Soc. N.S.W., 125, 2004 



225 



CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN PJVER 



Grant, T.R. and Whittington, R.J. (1991). The use of 

freeze-branding and implanted transponder tags 
as a permanent marking method for platypuses, 
Ornithorhynchus anatinus (Monotremata: 
Ornithorhynchidae). Australian Mammalogy 14: 
147-150. 

Gust, N., Handasyde, K. (1995). Seasonal variation in the 
ranging behaviour of the platypus 
{Ornithorhynchus anatinus) on the Goulburn 
River; Victoria. Australian Journal of Zoology 
43, 193-208. 

Hulbert, A.J. and Grant, T.R. (1983a). A seasonal study of 
body condition and water turnover in a free- 
ranging population of platypuses, 
Ornithorhynchus anatinus. Australian Journal 
of Zoology 31: 109-116. 

Hulbert, A.J. and Grant, T.R. (1983b). Thyroid hormone 
levels in the egg-laying monotreme, the 
platypus, Ornithorhynchus anatinus. General 
and Comparative Endocrinology 51: 401-405. 

Munday, B.L., Whittington, R.J. and Stewart, N.J. (1998). 
Disease conditions and subclinical infections of 
the platypus {Ornithorhynchus anatinus). 
Philosophical Transactions of the Royal Society 
London. Biological Sciences 353, 1093-1099. 

Rakick, R., Rakick, B., Cook, L. and Munks, S. (2001). 
Observations of a platypus foraging in the sea 
and hunting by a wedge-tailed eagle. Tasmanian 
Naturalist 123, 3-4. 

Serena, M. (1994). Use of time and space by platypus 
{Ornithorhynchus anatinus; Monotremata) 
along a Victorian stream. Journal of Zoology 
(London) 232, 117-131. 



Serena, M., Thomas, J.L., Williams, G.A., Officer, R.C.E. 
(1998). Use of stream and river habitats by the 
platypus, Ornithorhynchus anatinus, in an urban 
fringe habitat. Australian Journal of Zoology 
46, 267-282 

Temple-Smith, P.D. (1973). Seasonal breeding biology of 
the platypus, Ornithorhynchus anatinus Shaw 
1799, with special reference to the male. PhD 
Thesis. Australian National University: 
Canberra. 

Whittington, R.J. (1991). the survival of platypuses in 

captivity. Australian Veterinary Journal 68, 32- 
35. 

Whittington, R.J. and Grant, T.R. (1983). Haematology 
and blood chemistry of free-living platypuses, 
Ornithorhynchus anatinus 
(Shaw)(Monotremata: Ornithorhynchidae). 
Australian Journal of Zoology 31: 475-482. 

Whittington, R.J. and Grant, T.R. (1984). Haematology 
and blood chemistry of the conscious platypus, 
Ornithorhynchus anatinus 
(Shaw)(Monotremata: Ornithorhynchidae). 
Australian Journal of Zoology 32: 631-635. 

Whittington, R.J. and Grant, T.R. (1995). Haematological 
changes in the platypus {Ornithorhynchus 
anatinus) following capture. J. Wildlife Diseases 
31: 386-390. 

Whittington, R.J., Connolly, J.H., Obendorf, D.L., 

Emmins, J., Grant, T.R. and Handasyde, K.A. 
(2002). Serological responses against the 
pathogenic fungus Mucor amphibiorum in 
populations of platypus {Ornithorhynchus 
anatinus) with and without ulcerative mycotic 
dermatitis. Veterinary Microbiology 87, 59-71. 



226 



Proc. Linn. Soc. N.S.W., 125, 2004 



Breeding in a Free-ranging Population of Platypuses, 
Ornithorhynchus anatinus, in the Upper Shoalhaven River, New 

South Wales - a 27 Year Study 

T.R. Grant 1 , M. Griffiths 2 and P.D. Temple-Smith 3 

1 School of Biological, Earth and Environmental Sciences, University of NSW, Sydney 2052, 

t.grant@unsw.edu.au; 2 80 Dominion Circuit, Deakin, ACT, 2600 

department of Conservation and Research, Zoological Parks Board of Victoria and 

The University of Melbourne 



Grant, T.R., Griffiths, M. and Temple-Smith, P.D. (2004). Breeding in a free-ranging population of 
platypuses, Ornithorhynchus anatinus, in the upper Shoalhaven River, New South Wales - a 27 year 
study. Proceedings of the Linnean Society of New South Wales 125, 227-234. 

A total of 150 captures of lactating platypuses (97 individuals) were made over a period of 27 years in the 
study area. The proportion of lactating females from December samples ranged from 18 to 80% (mean 
43.4±17.7%; n = 21 breeding seasons). The percentage of juveniles in samples taken at the seasons when 
young were leaving the nesting burrows varied from 0-63% (mean 34.4±17.9%; n = 22 breeding seasons). 
Only 8.8% percent of captured juvenile females went on to breed in the area; one bred in its second breeding 
season after emergence but two others did not breed until at least their 4 th breeding season. Some females 
bred during at least 2-3 consecutive breeding seasons but others failed to breed in consecutive years. The 
percentages of females lactating in the months of September to April indicated a spread in the breeding 
season. Lactation in the wild was apparently shorter than reported in captivity, lasting more than 3 but less 
than 4 months. The majority of variation in breeding activity and recruitment could not be explained in 
terms of drought or observed riverine and riparian changes during the study. 

Manuscript received 2 September 2003, accepted for publication 8 January 2004. 

KEYWORDS: Breeding, Drought, Lactation, New South Wales, Ornithorhynchus anatinus, Platypus, 
Recruitment, Sedimentation 



INTRODUCTION within the study area, with sand slugs encroaching into 

many of the pools and considerable bank erosion 

Platypuses {Ornithorhynchus anatinus) mate occurring as a result of poor past and present riparian 

in late winter or early spring. Eggs are laid and the and catchment land management practices. On 

developing young are nourished on milk in the nesting completion of the initial studies early in the 1990s, the 

burrows for several months, after which juveniles leave investigation continued by sampling in December, 

these burrows, become independent and most disperse when females captured would be most likely to be 

from natal sites. There is a north-south cline in the lactating, and in February or March when juveniles 

timing of the breeding season, which begins earliest had left the nesting burrows but had not yet dispersed 

in north Queensland and latest in Tasmania (Temple- (see also Grant, 2004 this volume). 
Smith and Grant 2001). The current study was carried While the study has permitted general aspects 

out near the centre of this cline, on the southern of lactation to be further considered since the work of 

tablelands of New South Wales in the upper Grant and Griffiths (1992), it has also investigated the 

Shoalhaven River. It began with the investigation of effects of stream degradation and drought on the 

the nature of lactation and the composition of the milk platypus population in the upper Shoalhaven River, 

of the platypus (Griffiths etal. 1973; Grant etal. 1983; With regard to this latter aspect of the study, the 

Messer et al. 1983; Parodi and Griffiths 1983; Griffiths hypothesis being tested was that successful breeding, 

et al. 1984; Griffiths et al. 1985; Gibson et al. 1988; as indicated by females breeding and young being 

Teahan et al. 1991; Grant and Griffiths 1992; Joseph recruited to the population each year, would be 

and Griffiths 1992). However, in the mid-1980s there adversely affected by stream degradation and/or by 

was considerable change to the habitat of the platypus droughts. 



BREEDING IN FREE-RANGING PLATYPUSES 



MATERIALS AND METHODS 

Study Site 

The study area consisted of a series of 16 
pools in agricultural land, separated by riffle areas 
along 12.5 kilometres of the upper Shoalhaven River 
and 3.9 kilometres of an adjacent tributary stream, near 
Braidwood on the southern tablelands of New South 
Wales. A narrow discontinuous strip of riparian 
vegetation, consisting of both introduced and 
indigenous species of trees and shrubs, interrupted by 
numerous gaps, which were normally eroded as a result 
of access by sheep and/or cattle from the surrounding 
pasture land to the river. During the period of the study 
(late 1977 to early 2004), some of these pools suffered 
in-filling by sand slugs. For some pools, the effects on 
the habitat from sand in-filling was so severe that they 
were deleted from the sampling program. During the 
study period, three significant droughts occurred. 

Sampling Periods 

Two to four pools representative of the area 
(core area) were sampled during December, then again 
in February and/or March of 21 and 22 breeding 
seasons respectively over the 27 years of the study. 
Other pools within and outside these core area pools 
were sampled intermittently at various times during 
research in associated projects (Grant, 2004 this 
volume). 

Capture, Marking and Possessing 

Animals were captured using the unweighted 
"gill" net methods outlined in Grant and Carrick 
(1974). Until 1987, individuals were marked using 
stainless steel leg bands (Grant and Carrick 1974) but 
these were phased out after trials on the use of Passive 
Integrated Transponder (PIT) tags proved to be 
successful (Life Chip tags; Destron Fearing 
Corporation Scanner; Grant and Whittington 1991; 
Grant 2004; this volume). After removal from the nets, 
animals were weighed, measured and age and sex were 
determined (Temple-Smith 1973; Grant 2003 this 
volume). Females were injected intramuscularly with 
0.1-0.2 mL of synthetic oxytocin (1-2 International 
Units; Syntocinon, Novartis) to induce milk "let down" 
(Griffiths et al. 1972, 1973, 1984; Grant and Griffiths 



1992). In females that were lactating, milk could be 
expressed from the mammary gland, using gentle 
pressure along the flanks towards the areolae, 5 minutes 
after injection. 

Data Collection 

The percentage of lactating females captured 
in each December sample and the percentage of 
juveniles caught in relation to the total numbers of 
animals captured at each sampling in February and/or 
March were calculated. These provided indices of 
breeding and recruitment success for each breeding 
season. The timing and duration of lactation were 
determined from these data and from the capture and 
recapture of females in other pools of the study area. 



RESULTS 

Timing and duration of lactation 

During the 27 years of the study, captures of 
150 lactating platypuses were made. A total of 97 
individuals were lactating at least once during the study 
(Table 1). Only a single individual was found lactating 
in late September, with the highest proportions of 
lactating animals being captured in December and 
January. Sequential recaptures of three individuals 
within the same breeding season showed lactation in 
the field lasted at least 70-98 days (2.3-3.3 months) 
(Table 2). Other sequential data showed that 97% (30 
from 31) of females found lactating in December or 
January, had ceased lactation when recaptured in 
March. Of five individuals lactating in December, three 
were no longer lactating when recaptured in February 
(Table 2). 

Breeding ages of juvenile platypuses 

Of 137 female platypuses captured as 
juveniles, only 12 were later recaptured as breeding 
females in the study area (8.8%, Table 3). One of these 
individuals was lactating in its second breeding season 
after emergence from the nesting burrow, but three 
others did not breed until at least their 3 rd or 4 th breeding 
season. The individual (FJ222) which bred in the 
second breeding season (1983/84) failed to breed the 
following year (1984/85). This animal was not captured 





Sept. 


Oct. 


Nov. 


Dec. 


Jan 


Feb. 


Mar. 


Apr 


Total 
Lact. 


26 
1 


5 
1 


25 
7 


256 
106 


60 

24 


59 
10 


179 
1 


5 



% 


3.8 


20.0 


28.0 


41.3 


40.0 


17.6 


0.6 






Table 1. Numbers and percentages of individual female platypuses lactating in all samples in the upper 
Shoalhaven River study area 



228 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH 



Animal 



December 



January 



February 



March 



April 



Lactation 
duration 



FA015 


V 


- 


FA019 


S 


- 


FA046 


S 


- 


FA126 


S 


• 


FA133 


V 


- 


FA158 


V 


- 


FA158 


S 


- 


FA161 


V 


- 


FA161 


S 


- 


FA185 


V 


- 


FA185 


S 


-. 


FA185 


V 


- 


FA209 


S 


- 


FA212 


V 


- 


FA214 


S 


- 


FA276 


S 


- 


FA335 


S 


- 


FA368 


</ 


- 


FA368 


- 


• 


FA370 


V 


- 


FA391 


s 


- 


FA462 


s 


- 


FA514 


V 


- 


FA535 


</ 


■' - 


FA530 


- 


• 


FA547 


^ 


- 


FJ157 


s 


- 


FJ222 


</ 


- 


FJ248 


s 


- 


FJ248 


- 


V 


FJ272 


s 


- 


FJ272 




s 


FJ272 


s 


s 


FJ436 


s 


- 


FJ469 


V 


- 



X 



X 



X 


'- ■ 


-. 




• 


X 


' - 


>72days 


- 


X 






- 


^ 


X 


>98 days 


1- 


X 


- 




- 


- 


X 




- 


X 


' - 




- 


X 


..■'.■- 




- 


X 


- 




- 


X 


- 




- 


X 


■ - 




- 


X 


- 




- 


X 


- - 




- 


X 


- 




- 


X 






- 


X 


- 




- 


X 


- 




s 


X 


- ■ 


>70 days 


- 


X 


- 




- 


X 


- 




- 


X 


- 




- 


X 


- 




- 


X 


■ - 




Y 


X 


- 




A 


X 


_ " 




- 


X 


- 




- 


X 


- 




- 


X 


- 




- 


X 


- 




- 


X 


' - 




- 


X 


- 




- 


X 


- 





Table 2. Recaptures of lactating platypuses within given breeding seasons S Lactating; X Not 
lactating; - not recaptured. 



in the breeding season of the next year (1985/86) but 
was lactating again in the subsequent breeding season. 

Breeding success and recruitment 

Twenty-eight females were captured in 
successive breeding seasons. While some females were 
captured lactating in up to three consecutive breeding 
seasons, many failed to breed in consecutive seasons, 
with 39% not lactating in a season immediately 
following one in which they did breed. (Table 4). 

The mean percentage of lactating (breeding) 
animals in December of 21 breeding seasons in the 
core section of the study area was 43.4±17.7% but the 
numbers and proportions fluctuated considerably 
between breeding seasons from 80% down to 18% of 
the numbers of females captured (Figure 1). 



The data showed a general relationship 
between the numbers of juvenile platypuses captured 
in February and/or March in the core area of the study 
site and the total numbers of lactating animals captured 
in each of the breeding seasons sampled (Figure 1). In 
some years recruitment of juveniles was low after 
reasonable numbers of lactating females had been 
captured in December and in other years higher than 
expected recruitment levels were observed in February/ 
March after relatively low numbers of lactating females 
had been captured in the previous December. However, 
in general higher percentages of juveniles were 
captured at the end of breeding seasons when the 
percentage of lactating females sampled was also high 
(Figure 1). 



Proc. Linn. Soc. N.S.W., 125, 2004 



229 



BREEDING IN FREE-RANGING PLATYPUSES 



"i ST 

— ■ 

3 (V 

© © 

SO if -■ 

.. 3 — JT 



S ©" 

» 3 

© Q. 

3 2; 

p OTQ 

n 

© 

3 

a 

O" 

I-! 
ffi 
ffi 

a 
5' 

CTQ 

as 

re 

B9 
03 

o 

3 
85 

3? 

CD 

ft 



ffi IS 

a *< 

TO "2 

**• u< 

<v ffi 

ST © 

n 2, 

£ CTQ* 

3 65 
QTQ CJ 

3o 

2 T3 

3 if 

S.& 

ffi — • 

65 3 

•B «■ 

e B* 
f ffi 

ffi w 

^ S. 
TO 5* 

© 5. 

< 3* 

3 ~ 

3 cr 
o- — 
ffi as 

ZL S" 

^* -j 

5 "* 
© « 

c 3! 

g. 3 3 

3 D. tt 



3 
BO 

a* 

3 

»l 
O 



a* 
•-» 
re 
« 
a 

5' 

QTQ 

65 



65 



»ga 



I 

4- 

o* 

ffi 
ffi 

a 
5' 

CTQ 



ST ©- 
ffi i^i 

^. ft} 



5' '* 

If * 



r^ ^5 ^3 ^9 ^5 

c_i «-< e-i Sh e-t 

-E* 4i. «k Ji. N» 

s© ©\ w © -a 

On v© ©\ V© W 



W W W 



Tl •=) 



to to to to bO to K- 

-a f w w is> m en 
to wuo to -a o 



* * 

* * 



X! 



X. 



x n 

X ' 
X 

■ ■ X! 



>|\ H 



* a \S X 



lx>« X 

■ X! - S 

\ X! 



* X! 



s*s 



\ 



X ■ \ 



* X < . . 

* s ^ 

X X i i 



■ 




X 


\ 


n 


\ 



X 



■ I I 



>tx X X \ 



• n\ 



n 



n 



\ 



X. 



K x 



n\ 



\ 



\ 



\ 



n 



n\ 



n 



oo 2S 



vo ^© 



to 



230 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH 



g ITS 

3. ©, 2 

QTQ 

2 „ 3 

g " tB 

c* 2 re 
» a*E. 

& C P 
65 5 J? 

f! S fl 

cr cro c 
2 jt re 

a s 5' 
5« J- © 

re ^ ^ 

-* re 2 

?3| 

q, i re 
*~to n 
& re _. 

£ a « 

1ST" 

re N ^ 

_ 65 © 

aT re a 

I £ 8 
»are 

<-►■ 3 C 

a'5fi 3- 
TO - * re 

re o « 
H ~ 2 

£ O 3 

was 

^-^ 55 re 

• 3 as 

TO £ 

-• © 

•>s a 

re _ 

g B 
to a 
s=" ST 



'TC] *T!l *Ti *Ti *Tj *T] 'Tj 

> J> > £ > >£ 
ui w w io h h ffi 






*Ti *Tl "Tl *Tl *T} 
>>>>> 

o\ ui *. u> to 

H-» 00 -O. U> ON 



XX 



* K> U KJ 

# ?^ W K^ 



X S \ 



xxxx 
.*xx ■ 
XI x x •>» x ■ 
X ■ • X \x \x 



\ 



\xx \ \x 



\ 



X 

X 

XXX 

■*xxx \ 

X ■ X •* X ■< 

x \x \-* \ 

' XX ■ X • 

I I I I I I 

. .* \ . \ . 
V' ■ ■ \ ■ 



\ 



\ 

\ 
\ 

X 



X ■ X 
X V* 

\ \x 

^x 
• \ 
\ s 

X ■ 



X 

\ 
\ 



s 



X 



s 



\ 



•■j— s •< X 

\X ' \x 
WW 

\ \xx 



\ 



\ 

X 



to >-> 
■J^ no 


r. 

B 

E. 


' 




1 


\Q 00 


X \ 


0» ^ 


S ' 


00 00 

h-5 


■ \ 


OS ao 


■ \ 


00 2? 


1 .^ 


00 pS 


■ \ 


OS os 




os p£ 




oe 2? 




06 06 
OS5J 




os 2S 

V© 06 




>0 06 

© ^ 








Sfe- 




\e no 




VO 'O 




\a no 




ve n© 

ON S5 




<I?N 




SO N© 




ve n© 

v© ss 




o ^ 








o o 








2| 



Proc. Linn. Soc. N.S.W., 125, 2004 



231 



BREEDING IN FREE-RANGING PLATYPUSES 



GO 



04 

c 



« 



100 i 
90 
80 
70 - 
60 - 
50 
40 
30 
20 

io H 




I 






I 









Breeding Seasons 

Figure 1. Percentages of lactating females captured in relation to total adult females caught in each 
December sample (n = 21) and the percentages of juveniles in each February and/or March sample (n 
= 22) in the core study area. Open bars = lactating females (5 seasons not sampled); solid bars = 
juveniles (4 seasons not sampled; 1988/89 and 1991/92 seasons no juveniles were caught). 



DISCUSSION 

The low overall percentages of lactating 
animals in samples caught in September (3.8%), 
October (20%), November (28%) and in February 
(17.6%), March (0.6%), April (0%), contrasted with 
higher percentages in December (41.3%) and January 
(40%). This indicated a spread in the breeding season. 
However, it appears that the majority of animals were 
breeding around the same time, with a few individuals 
breeding earlier (eg. one already lactating by the end 
of September) and a few later (eg. one still lactating in 
March; Table 1). 

The sequential recaptures of lactating females 
within the same breeding seasons provided evidence 
that lactation in the wild can last at least 98 days (3.3 
months) but is unlikely to exceed four months. This 
suggestion is supported by the distribution of lactating 
females in the various months (Table 1), combined 
with the observation that all but one of 31 females 
lactating when captured in December or January (97%) 
had ceased lactation on their subsequent recapture in 
March (Table 2). 



Nestlings in the wild may be weaned more 
rapidly than those bred in captivity. Lactation in captive 
animals has been reported to continue as long as 145 
days (4.8 months; Holland and Jackson 2002; 
Healesville Sanctuary and Taronga Zoo, unpublished) 
and requires lactating females to consume up to 100% 
of their body weight in food during peak lactation 
(Holland and Jackson 2002). It may be that the young 
are weaned more quickly in the wild depending on the 
local availability of macroinvertebrate food items 
(Faragher et al. 1979) for the breeding females. 
Certainly in some years of this study, lactating females 
were in poorer body condition, based on observations 
of tail fat reserves (Temple-Smith 1973; Grant and 
Carrick 1978) and general body condition. 
Interestingly, both the lactating females and the 
captured juvenile animals in the 2002/03 breeding 
season, at the end of a very severe drought, were judged 
to still be in good body condition. However, four 
lactating females captured at the beginning of January 
2004 appeared to be in poorer condition, despite the 
river flows being an improvement on those of the 
previous breeding season, when surface flows in the 



232 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH 



study area stopped for several weeks (pers. comm. 
from local residents). 

Recapture rates of juvenile platypuses during 
the various studies in the area was quite low. Thirty- 
two percent of female platypuses first captured as 
juveniles were recaptured compared to only 14% for 
males (Grant 2004; this volume). In spite of this, only 
12 female juveniles were recruited into the breeding 
population. 

Although the breeding season was 
predictable, the breeding of individuals in any one 
season was much less predictable, with varying 
numbers of non-breeding females in any sample, 
individuals not breeding until later in life and breeding 
animals failing to breed in consecutive seasons (Table 
4). Similar observations have been made with 
platypuses in captive conditions, where no individuals 
have so far bred in successive breeding seasons 
(Holland and Jackson 2002; Healesville Sanctuary and 
Taronga Zoo, unpublished). Temple-Smith and Grant 
(2001) have speculated whether resource availability, 
social organisation or genetic factors are involved in 
this uncertain breeding in the species but little is known 
of any of these aspects of platypus biology. 

A decline was expected in the number of 
platypuses breeding and/or the number of juveniles 
recruited to the population after the mid 1980s, when 
sand slugs began to reduce the pools available for 
foraging and the provision of refuge areas during 
drought. Surprisingly, no such overall trend occurred 
in either numbers breeding or in recruitment data and 
there is no ready explanation for the considerable 
variation in the numbers breeding between the seasons 
covered by the study. Such variations must be 
attributable to more subtle changes occurring in the 
environment and/or to unexplained sampling effects. 

Both breeding success and recruitment fell 
sharply in the 1982/83 breeding season, corresponding 
to the end of a long and severe drought, which lasted 
from October 1978-February 1983. The effects of the 
drought provided an explanation for the observation 
that three females lactating in the 1981/82 season did 
not breed in the 1982/83 season. However, there were 
no similar trends recorded in the 1993-95 or 2001-03 
droughts, although during the latter, lactation and 
recruitment percentages were slightly below the mean 
values for each (Fig. 1). As discussed above, all 
lactating females and juveniles captured in the 2002/ 
03 sampling were considered to be in good body 
condition in spite of the severe drought conditions 
which existed at the time. 

During the 1988/89 and 1991/92 breeding 
seasons no juvenile platypuses were captured. There 
is no obvious reason for the observed lack of 
recruitment in 1988/89, but two local over-bank flood 



events in late December/early January (pers. comm. 
from local residents) of 1991/92 may have drowned 
many nestlings confined to burrows during that season. 
This would explain the failure to capture juveniles in 
a year when the percentage of lactating females in the 
previous December had been slightly higher the mean 
value (Fig. 1). 

Prior to enactment of legislation protecting 
platypuses in all states of Australia (1892 in Victoria 
to 1912 in South Australia; Grant and Denny 1991), 
thousands were hunted for their fur. Their numbers 
are reputed to have declined dramatically, although 
rebounding since protection has been enforced (Grant 
and Denny 1991 ; Grant and Temple-Smith 1978). The 
species is currently listed as protected, but is either 
regarded as 'common' or not threatened, in all states 
(except for South Australia, where it is probably now 
extinct, except for an introduced population on 
Kangaroo Island). In spite of this, there is concern at 
the fragmentation of populations in some river systems 
and in small local populations as a result of habitat 
degradation, illegal and recreational fishing and 
encroaching effects of urban and regional development 
(Grant and Temple-Smith 2003). While this study 
demonstrates that the species has continued to survive 
and reproduce in the upper Shoalhaven River in spite 
of considerable riparian and riverine degradation, the 
effects of drought and the combination of both of these 
perturbations, further investigation leading to a 
complete understanding of the factors determining the 
uncertain breeding in the species is critical to its 
conservation. Many questions regarding the population 
biology and reproduction of Ornithorhynchus anatinus 
still remain unanswered but the long-term studies 
reported here and in Grant (2004, this volume) have 
gone some way to providing a greater understanding 
of some aspects of the species' field biology, which 
could not have been achieved by a study of shorter 
duration. 

ACKNOWLEDGMENTS 

Merv Griffiths, a friend, colleague and our co- 
author died on 06 May 2003. The many other friends and 
colleagues who were instrumental in the success of field 
work, in often severely inclement conditions, over the years 
are too numerous to name individually but the Heath family, 
Paul Anink, Marie-Loiuse Lissone, David Read and Gina 
Grant deserve special mention. All are gratefully 
acknowledged. Some of the work reported was done while 
in receipt of funding from the Environment Australia (then 
Australian National Parks and Wildlife Service) and the 
Australian Research Council (then Australian Research 
Grants Committee). The late Athol MacDonald and the 
Izzard and Laurie families are acknowledged for their 
permission to access the river and creek on the properties 
managed or belonging to them, and for their friendship and 



Proc. Linn. Soc. N.S.W., 125, 2004 



233 



BREEDING IN FREE-RANGING PLATYPUSES 



assistance in various aspects of the field work. The work 
was carried out under NSW National Parks and Wildlife 
Service Scientific Investigations Licence A 184, New South 
Wales Fisheries Scientific Research Permit F84/1245 and 
University of New South Wales Animal Care and Ethics 
Approvals 94/91, 97/46 and 00/45. 



REFERENCES 

Faragher, R.A., Grant, T.R. and Carrick, F.N. (1979). 
Food of the platypus, Ornithorhynchus 
. anatinus, with notes on the food of the brown 
trout, Salmo trutta, in the Shoalhaven River, 
New South Wales. Australian Journal of 
Ecology 4: 171-179. 

Gibson, R.A., Neumann, M., Grant, T.R. and Griffiths, M. 
(1988). Fatty acids of the milk and food of the 
platypus {Ornithorhynchus anatinus). Lipids 23, 
377-379. 

Grant, T.R. (2004). Captures, capture mortality, age and 
sex ratios of platypuses, Ornithorhynchus 
anatinus, during studies over 30 years in the 
upper Shoalhaven River in New South Wales. 
Proceedings of the Linnean Society of New 
South Wales 125, 217-226. 

Grant, T.R. and Carrick, F.N. (1974). Capture and 
marking of the platypus, Ornithorhynchus 
anatinus, in the wild. Australian Zoologist 18: 
133-135. 

Grant, T.R. and Carrick, F.N. (1978). Some aspects of the 
ecology of the platypus, Ornithorhynchus 
anatinus in the upper Shoalhaven River, New 
South Wales. Australian Zoologist 20: 181-199. 

Grant, T.R. and Denny, M.J.S. (1991). Historical and 
Current Distribution of the Platypus in 
Australia, with Guidelines for the Management 
and Conservation of the Species. Unpublished 
Report to Australian National Parks and 
Wildlife Service by Mt. King Ecological 
Surveys. 

Grant, T.R., Griffiths, M. and Leckie, R.M.C. 1983. 
Aspects of lactation in the platypus, 
Ornithorhynchus anatinus (Monotremata), in 
waters of eastern New South Wales. Australian 
Journal of Zoology 31, 881-889. 

Grant, T.R. and Griffiths, M. (1992). Aspects of lactation 
and determination of sex ratios and longevity in 
a free-ranging population of platypuses, 
Ornithorhynchus anatinus, in the Shoalhaven 
River, New South Wales. In 'Platypus and 
Echidnas'. (Ed ML.Augee). pp. 80-89. (Royal 
Zoological Society of NSW: Sydney). 

Grant, T.R. and Temple-Smith, P.D. (1998). Field biology 
of the platypus (Ornithorhynchus anatinus) - 
historical and current perspectives. Transactions 
Royal Society London Series B 353, 1081-1091. 

Grant, T.R. and Temple-Smith, P.D. (2003). Conservation 
of the platypus, Ornithorhynchus anatinus: 
Threats and challenges. Aquatic Health and 
Management 6, 1-15. 



Grant, T.R. and Whittington, R.J. (1991). The use of 

freeze-branding and implanted transponder tags 
as a permanent marking method for platypuses, 
Ornithorhynchus anatinus (Monotremata: 
Ornithorhynchidae). Australian Mammalogy 14: 
147-150. 

Griffiths, M., Elliott, M.A., Leckie, R.M.C. and Schoefl, 
G.I. (1973). Observations on the comparative 
anatomy and ultrastructure of mammary glands 
and on the fatty acids of the triglycerides in 
platypus and echidna milk fats. Journal of 
Zoology [London] 169, 255-275. 

Griffiths, M., Green, B., Leckie, R.M.C, Messer, M. and 
Newgrain, K.W. (1984). Constituents of 
platypus and echidna milk with particular 
reference to the fatty acid complement of the 
triglycerides. Australian Journal of Biological 
Science 37, 323-329. 

Griffiths, M., Mcintosh, D.L. and Leckie, R.M.C. (1972). 
The mammary glands of the red kangaroo with 
observations on the fatty acid components of the 
milk triglycerides. Journal of Zoology [London] 
166, 265-275. 

Griffiths, M., McKenzie, H.A., Shaw, D.C. and Teahan, 
C.G. (1985). Monotreme milk proteins: echidna 
and platypus "whey" proteins. Proceedings of 
the Australian Biochemical Society. 17, 25. 

Holland, N. and Jackson, S.M. (2002). Reproductive 
behaviour and food consumption associated 
with captive breeding of platypus 
(Ornithorhynchus anatinus). Journal of Zoology 
[London] 256, 279-288. 

Joseph, R and Griffiths, M. (1992). Whey proteins in early 
and late milks of monotremes (Monotremata; 
Tachyglossidae, Ornithorhynchidae) and of the 
tammar wallaby (Macropus 
ewgenz7)(Marsupialia; Macropodidae). 
Australian Mammalogy 15, 125-127. 

Messer, M., Gadiel, P.A., Ralston, G.B. and Griffiths, M. 
(1983). Carbohydrates of the milk of the 
platypus (Ornithorhynchus anatinus). 
Australian Journal of Biological Science. 36, 
129-138. 

Parodi, P.W. and Griffiths, M. (1983). A comparison of 
the positional distribution of fatty acids in milk 
triglycerides of the extant monotremes platypus 
(Ornithorhynchus anatinus) and echidna 
(Tachyglossus aculeatus). Lipids 18, 845-847. 

Teahan, C.G., McKenzie, H.A. and Griffiths, M. (1991). 
Some monotreme milk "whey" and blood 
proteins. Comparative Biochemistry and 
Physiology. 91B, 99-118. 

Temple-Smith, P.D. (1973). Seasonal breeding biology of 
the platypus, Ornithorhynchus anatinus (Shaw 
1799) with special reference to the male. PhD 
thesis, Australian National University, Canberra. 

Temple-Smith, P.D. and Grant, T.R. (2001). Uncertain 
breeding: A short history of reproduction in 
monotremes. Reproduction, Fertility and 
Development 13, 487-497. 



234 



Proc. Linn. Soc. N.S.W., 125, 2004 



Depth and Substrate Selection by Platypuses, Ornithorhynchus 
anatinus, in the Lower Hastings River, New South Wales 

Tom Grant 

School of Biological, Earth and Environmental Sciences, University of New South Wales, NSW 2052 

t.grant@unsw.edu.au 



Grant, T. (2004). Depth and substrate selection by platypuses, Ornithorhynchus anatinus, in the lower 
Hastings River, New South Wales. Proceedings of the Linnean Society of New South Wales 125, 235- 
241. 

Platypuses were observed foraging most frequently in water >1 metre in depth during normal (91.3%) and 
drought (82.1%) conditions. Mean water depth in the study pools was 1.08±0.66 and 0.86±0.61 metres 
during normal and drought conditions respectively. The distribution of depths in the study area was 
significantly different from the distribution of depths where platypuses were observed during normal (Chi 2 
= 90.2; p < 0.01) and drought conditions (Chi 2 = 37.35; p < 0.01). Platypuses were apparently not simply 
utilising depths in relation to their occurrence but preferring to forage in water deeper than 1 .5 metres and 
avoided depths < 1 metre. Overall distribution in numbers of platypuses observed foraging over different 
benthic substrate types was not significantly different (Chi 2 = 12.9; p > 0.05) from the distribution of these 
substrate categories in the study area. However, when the substrates were considered separately, significant 
preference was shown for cobbled substrate (Chi 2 = 18.4; p < 0.01) and avoidance of gravel (Chi 2 = 9.7; p 
< 0.01). These observations have implications for catchment, stream and riparian management, where activities 
leading to sedimentation and reduced flushing flows may reduce depths and/or alter the distribution of 
preferred foraging substrates. 

Manuscript received 2 September 2003, accepted for publication 7 January 2004. 

KEYWORDS: depth, foraging, Hastings River, Ornithorynchus anatinus, platypus, substrate. 

of stream use and management activities. As part of 

INTRODUCTION the monitoring and detection of possible environmental 

effects of the Hastings District Water Supply 

During foraging in the wild, platypuses dive Augmentation Scheme, the utilisation of depth and 

to feed almost exclusively on small benthic invertebrate substrate categories by foraging platypuses was 

animals (Faragher et al. 1979; Grant 1982), which are investigated, 
normally unevenly and often sparsely distributed in a 
variety of substrates and depth zones (Boulton and 

Brock 1999; Elliott 1977; Young 2001). The platypus METHODS 
is small, has a high metabolic demand to regulate its 

body temperature in water and has an estimated Study area 

maximum aerobic capacity for diving of only 40-60 The study was undertaken in two separate 1.5 

seconds (Bethge 2002; Bethge et al. 2001; Evans et kilometre sections of the lower Hastings River near 

al. 1994; Grant and Dawson, 1978). Consequently its Wauchope in New South Wales. Immediately above a 

foraging is restricted to relatively shallow depths and lar ge riffle separating the riverine section from the 

the species is seldom reported occurring in deep lakes upper estuary tidal influence, the study area consisted 

or impoundments (Bryant 1993; Grant 1991; McLeod f a ser ies of pools, riffles and runs, with the banks 

1993; Ellem et al. 1998; Ellem and McLeod 1998). predominantly consisting of earth consolidated by the 

The current study reports on observations of depths of roots of riparian vegetation, but with a number of 

diving and foraging over different substrates by gravel/cobble bars and sections of bedrock also present 

platypuses in a coastal river in New South Wales during . Predominantly surrounded by agricultural land, 

drought and normal flow conditions. Grant and Bishop especially pastures for dairy cattle, much of the stream 

(1998) discussed the importance of the measurement bank supported a narrow strip of vegetation consisting 

of physical habitat variables associated with platypus f river oaks (Casuarina sp), rainforest species (e.g. 

occurrence as a means of assessing possible impacts Waterhousea floribunda, Ficus coronata) and 



DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES 



introduced weed species (e.g. willows, Salix sp; 
Lantana camara; privet, Ligustrum spp, wild tobacco, 
Solarium mauritianum). A range of macrophyte species 
also occurred in the stream (especially Myriophyllum 
verrucosum), although these were reduced to low 
incidences after several flood events. The aquatic grass, 
Potamophila parviflora, was also common along 
several sections of bank and occurred in island clumps 
within several sections of the stream. 

Sampling 

The 3 kilometres of river surveyed consisted 
of four riffle areas and five pools. Each section was 
surveyed in both directions during the two hours prior 
to darkness and immediately after first light in winter 
(late May to early July) and spring (September to 
October) over six years from 1998-2003 (88 
longitudinal transects x 2 river sections = 176 
longitudinal transects; i.e. the whole 3 kilometres was 
surveyed 88 times). During 1998-2000 the same 
number of longitudinal transects (16) was surveyed in 
both winter and spring but from 2001 to July 2003 
fewer were surveyed in winter (8) and more in spring 
(24), as lower numbers of platypuses were normally 
observed during the winter period. Depth and the 
predominant substrate type were recorded at the point 
where each platypus was first seen foraging. It should 
be noted that visibility, due to turbidity and/or poor 
light conditions, often meant that a determination of 
substrate could not be made at all of these points. 
During the 2001 and 2002 sampling periods, visibility 
was so low (probably due to the high abundance of 
phytoplankton) that the substrate could be observed 
only in few instances where platypuses were foraging. 

Physical habitat analysis 

The stream was paced out into 60 x 50 metre 
sections (3 km) and marked at each point with brightly- 
coloured flagging tape. At each of these points depth 
measurements were made at both edges (approximately 
2 metres from bank) and in the middle of the river 
(using a weighted line or the kayak paddle graduated 
in 25 cm units). These depth measurements (n = 174) 
provided a measure of the distribution of depth 
categories in the study area (Figs 1 and 2). The 
occurrence of benthic substrates (mud, sand, gravel, 
cobble and bedrock) was scored on a scale of 1-5 (using 
the following estimated percentage cover of each 
substrate type; = 0%; 1 = 0-5%; 2 = 5-25%; 3 = 25- 
50%; 4 = 50-75%; 5 = >75%) along three transects 
parallel to the stream bank between corresponding 
depth measurement points at the beginning and end of 
each 50 metre section. Substrates along these transects 
were not homogeneous, often with some of each type 
within a single transect. However, the predominant 



substrate types (score of 4 or 5; i.e. >50% estimated 
coverage) for each transect (n = 174) were used as a 
measure of the distribution of the occurrence of 
substrates (Fig. 3). Data on depths and substrate 
distribution were collected once during July 2000 but 
depth measurements were repeated in October 2002 
when the river was under severe drought conditions 
and was barely flowing. 

Data analysis 

The null hypothesis being tested was that the 
occurrence of platypuses across depth and substrate 
categories was the same as the occurrence of these 
categories in the study area. The overall distribution 
of numbers of platypuses observed foraging within the 
various depth and substrate categories and the recorded 
numbers of occurrence of these physical attributes in 
the stream (n = 174 samples), were compared using 
Chi 2 analysis (Statistica, StatSoft Inc.) with expected 
values calculated using 2x5 contingency tables (Bailey 
1969). Comparisons between numbers of platypuses 
observed foraging at specific depths or on specific 
substrates compared to those not foraging at these 
specific depths or substrates (i.e. all other depths or 
substrate categories) were made using Chi 2 for 2 x 2 
contingency tables. Comparisons between drought and 
non-drought measurements of depth were made using 
Student's t-tests for unpaired samples (Bailey 1969; 
Statistica; StatSoft Inc). Indices of selection/avoidance 
(Response Index) of depth or substrate categories by 
platypuses were calculated as: 
Response Index = 

% Occurrence of platypuses in a 

depth or substrate category 
% Occurrence of that depth or 
substrate category in the stream 
An index of greater than unity suggested a selection 
response and less than one an avoidance response to a 
depth or substrate category. All means given are ± 
Standard Deviation. 



RESULTS 

Depth selection 

The mean depths of the stream during non- 
drought and drought were significantly different, being 
1 .08±0.66 and 0.86±0.61 metres respectively (t = 3.48; 
p < 0.001). The maximum water depth recorded by 
these transect-based measurements was 2.75 metres 
but platypuses were recorded foraging at depths of up 
to 3.2 metres and the maximum depth recorded 
opportunistically (not along transects or at foraging 
sites) was just under 4 metres. 

Figure 1 shows the numbers of observations 



236 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT 




0.0-0.5 0.6-1.0 1.1-1.5 1.5-2.0 

Depth Category (metres) 



>2.0 



Figure 1. Percent occurrence of platypuses observed 
foraging in various depth categories (n = 127 
observations; white bars) and the percentage of 
occurrence of these depth categories (n = 174 
observations; black bars) in the Hastings River 
study area during non-drought conditions. 



60 



Jl 



I 



0.0-0.5 0.6-1.0 1.1-1.5 1.5-2.0 >2.0 

Depth Category (metres) 

Figure 2. Percent occurrence of platypuses observed 
foraging in various depth categories (n = 28 
observations; white bars) and the percentage of 
occurrence of these depth categories (n = 174 
observations; black bars) in the Hastings River 
study area during drought conditions. 



of platypuses foraging in the various depth categories 
during a range of non-drought conditions. These data 
show that 91.3% of the platypuses (total n = 127) were 
observed foraging in depths greater than 1 metre, 
despite this depth category occurring in only 39.1% 
of the study area. The distribution of numbers of 
platypuses foraging within the depth categories and 
the recorded numbers of occurrence of these categories 
were significantly different (Chi 2 = 90.2; p < 0.01). 

Platypuses showed significant preferences for 
water deeper than 1.5 metre (Response Indices 3.1- 




Mud 



Sand Gravel Cobbles Bedrock 

Substrate Category 



Figure 3. Percent occurrence of platypuses observed 
foraging in various substrate categories (n = 56 
observations; white bars) and the percentage of 
occurrence of these substrate categories (n = 174 
observations; black bars) in the Hastings River 
study area during non-drought and drought 
conditions. 

3.2) and avoidance of depths of less than 1 metre 
(Response Indices 0.2-0.7). Foraging within the 1.1- 
1.5 metre category showed no significant preference 
or avoidance by platypuses (Table la). 

During severe drought conditions (July and 
October 2002) there was a significant difference 
between the distribution of foraging platypuses (n = 
28) and the distribution of recorded depth categories 
(Chi 2 = 37.35; p = < 0.01; Fig. 2). Response Indices 
showed a similar, but apparently more marked pattern 
towards preference for depths > 1 metre (Response 
Indices 2.6-6.2) and avoidance of shallower depths 
(Response Indices from 0-0.7). Considering the small 
sample sizes of platypuses foraging in specific depth 
categories (n = 0-10) no Chi 2 analyses were attempted 
on these data collected during the drought. 

Substrate selection 

The numbers of platypuses observed foraging 
on particular benthic substrate types (n = 56) are shown 
in Figure 3. The overall distribution of numbers of 
platypuses foraging over the various substrates was 
not significantly different from the distribution of these 
substrates (Chi 2 = 12.9; p > 0.05). However, only 
26.8% of platypuses were found foraging over gravel 
substrates, while this substrate type was the most 
abundant in the study area (50.9%). While cobbles 
made up only 12.7% of the available substrate, 28.6% 
of the platypuses observed were foraging over this 
substrate. Response Indices of 0.5 and 2.3 respectively 



Proc. Linn. Soc. N.S.W., 125, 2004 



237 



DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES 



Table 1. Chi 2 and probability values (2x2 contingency tables; Statistics; StatSoft Inc.) for comparisons 
of: 



a. platypuses foraging at specific depths and those not foraging at each of these depths 


Depth Category 0.0-0.5 m 0.6-1.0 m 1.1-1.5 m 1.6-2.0 m 


>2 m 


Chi 2 27.1 38.8 4.3 18.8 

<0.01* <0.01* >0.05 <0.01* 


4.3 
<0.01* 


b. platypuses foraging on specific substrates and those not foraging on each of these substrates 


Substrate Category Mud Sand Gravel Cobbles 


Rock 


Chi 2 0.01 0.96 9.70 18.70 
>0.05 >0.05 <0.01* <0.01* 


0.26 
>0.05 



* i 



indicates statistical significance. 



for gravel and cobbles, suggested avoidance of the 
former and preference for the latter. Individual 
comparisons between platypuses foraging over specific 
substrates compared to all other substrates showed 
significant differences from expected for gravel and 
cobbles but not for the other substrate types (Table 
lb). 



DISCUSSION 

The study suggested that platypuses observed 
in the early morning and late afternoon/evening were 
selecting the deeper sections of their habitat, with 
91 .3% of the observed individuals foraging in water > 
1 metre in depth and 33. 1 % foraging in water of greater 
than 2 metres, which constituted only 39. 1 % and 10.3% 
of the area respectively during normal flow conditions. 
Even during the severe drought conditions, 82.1% of 
platypuses were still observed foraging in water deeper 
than 1 metre, despite the fact that the proportion of 
recorded depths > 1 metre had decreased by 11.5%. 
During the drought observations, 48.3% of the area 
had a depth of 0.0-0.5 metres but no platypuses were 
observed forging in this depth category. Thus, 
platypuses appeared to show preference for foraging 
in deeper areas and an avoidance of shallower depths 
within the area during the study in both drought and 
non-drought conditions. 

These observations were similar to those 
reported for a study in a small alpine lake in Tasmania 
(Lake Lea). While reporting a maximum dive depth 
of 8.77 metres, Bethge (2002) and Bethge et al. (2003) 
found that the majority of dives recorded for platypuses 
fitted with data loggers were to depths of less than 
three metres (98% of dives), with a mean diving depth 
of 1.28 metres. These workers also found a large 



proportion of the foraging dives were to depths of less 
than one metre (48%), although, during winter (when 
the lake level was higher than in summer), most dives 
were to depths greater than 1 metre. Platypuses were 
monitored foraging in the same area, rather than 
moving to the shallower parts of the lake during the 
winter, suggesting that foraging was determined by 
factors other than depth preference, possibly including 
substrate type and/or availability of benthic food 
species. However, the workers in this study did not 
report on these possibilities. 

Rohweder (1992), Bryant (1993) and 
McLeod (1993) also reported platypuses foraging in 
water less than 5 metres in depth. While Ellem et al. 
(1998) found increasing depth of pools (up to 2 metres) 
to be positively related to the observed presence of 
platypuses in 36 pools on the Macquarie River system 
in the Bathurst area of the central tablelands of New 
South Wales, Ellem and McLeod (1998) found radio- 
tacked platypuses using shallower parts of some 
sections of a weir pool in the Duckmaloi River near 
Oberon in New South Wales. 

Bethge (2002) also reported platypuses 
foraging in deeper areas and spending less time on the 
surface of the water in Lake Lea during daylight hours 
than at night. He speculated that these behavioural 
changes may have been related to avoidance of 
predators. Little is known regarding predation by 
indigenous predators which could take foraging 
platypuses from the water. Predation by the introduced 
red fox (Vulpes vulpes) on platypuses moving through 
or foraging in shallow water, such as riffle areas, has 
been reported (Serena 1994; Grant 1993; Anon. 2002) 
and the species has been included as a possible food 
item of wedge-tailed eagles (Aquilct audax){ Rakick 
et al. 2001; Marchant and Higgins 1993). 

Several white-breasted sea eagles (Haliaeetus 



238 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT 



leucogaster) and several ospreys (Pandion haliaetus) 
were observed at the Hastings River study site. Both 
of these species were seen taking fish from the surface 
of the water. Although a grey goshawk (Accipiter 
novaehollandiae) has been reported attacking a 
juvenile platypus on land (Richards 1986), it seems 
unlikely that either this species or the osprey would 
be large enough to take even a juvenile platypus (which 
are around 65-70% adult weight when they first leave 
the nesting burrows; Grant and Temple-Smith 1998) 
from the water. It is possible however, that the sea 
eagle may represent a potential predator of the 
platypus. During the study a sea eagle was observed 
retrieving a large dead Australian bass {Macquaria 
novemaculeata) of 485 mm in length, and estimated 
to weigh 2.5 kg (Harris 1987), from the bottom of a 
pool. The eagle was unable to fly with the fish and 
dragged it to a nearby gravel bar, where part of the 
flesh was eaten before darkness fell. Soon after first 
light the following morning the eagle was seen carrying 
off the remaining carcass of the bass. Interestingly, 
the platypus does not seem to have been recorded as a 
food item of this species of large eagle (Marchant and 
Higgins 1993; Olsen 1999). 

Substrate selection 

Higher invertebrate productivity is often 
associated with areas where logs, roots and vegetation 
provide a range of habitats for an array of types of 
benthic invertebrate species and coarse substrates 
(gravel, cobbles, rocks) provide fixed habitat, rather 
than a shifting substrate, such as sand and fine sediment 
(Young 2001; Boulton and Brock 1999; Smith and 
Pollard 1998). Data in the study were restricted by a 
small sample size due to the inability to observe the 
type of substrate over which platypuses were seen 
foraging in times of high turbidity and/or poor light 
conditions. However, there was some indication that 
platypuses were avoiding sections of the study area 
which consisted mainly of mud or sand (Fig 3) but 
this was not statistically significant (Table lb). There 
appeared to be marked avoidance of gravel (the most 
abundant substrate; 50.9% of the area) and preference 
shown for areas where cobbles were the predominant 
substrate (12.7% of the area). Both of these trends were 
statistically significant (Table lb). 

The complexity of benthic habitat has been 
previously identified as being positively related to the 
occurrence of platypuses (Rohweder 1992) and Serena 
et al. (2001) found a positive relationship between 
numbers of radio-tracked platypuses and the 
occurrence of coarser substrates, including gravel, 
pebbles, cobbles, large rocks and coarse particulate 
organic matter. These observations may be related to 
the distribution of benthic food organisms but this was 



not investigated in the present study. No explanation 
of the apparent avoidance of gravel substrate in the 
present study is suggested as the species has been 
observed by the author foraging on gravel substrates 
in other areas. 

Implications for stream management 

The development of adaptive management 
strategies for streams, particularly with regard to water 
extraction and the operation of impoundments, should 
consider flows which maintain pool depth and benthic 
habitat diversity by preventing the accumulation of 
sand and fine sediments. The removal of riparian 
vegetation, erosion as a result of unrestricted stock 
access to stream banks and poor catchment 
management practices have also resulted in the infilling 
of pools by sand 'slugs' in many streams in eastern 
Australia (Brooks and Brierley 1996; Boulton and 
Brock 1999; Brierley et al. 1999; Grant et al. 2003). 

Grant and Bishop (1998) encouraged the use 
of physical habitat analysis, considering broad habitat 
variables normally associated with platypus 
occurrence, in any attempts to monitor and/or predict 
effects of human activities impinging on streams and 
their catchments. More recently a habitat simulation 
model was used by Davies and Cook (200 1 ) to generate 
weighted useable habitat area estimates for the platypus 
at various proposed discharge regimes in a regulated 
river in Tasmania. This model used more specific 
habitat requirements of the species in terms of depth, 
velocity and substrate, calculating habitat preference 
curves based on available information from the 
literature and from experts in the field. These authors 
observed that: "Platypus[es] are known to feed in very 
shallow water and up to ca 1-3 m" and "foraging is 
optimal at depths of < 2 m" and "platypus[es] actively 
feed in silt, sands, finer gravel substrates, and are 
known to forage on coarse gravel to smaller cobble 
substrate. Feeding activity is not deemed to be efficient 
or to frequently occur on coarse cobble, boulder or 
bedrock substrates" 

While the important modelling work of 
Davies and Cook (2001) drew upon the information 
available to the authors at the time, the data from the 
current study and that from Serena et al. (2001) do not 
totally support the information used to generate their 
habitat preference curves for the species in mainland 
sites. It is vitally important that studies seeking to 
predict possible impacts of human activities on the 
platypus (or any other species) must consider the 
widest range and the most currently available 
information on which to base assumptions. 

Too often, assessment of possible 
environmental impact is based on 'conventional 
wisdom' which may be enshrined in publications 



Proc. Linn. Soc. N.S.W., 125, 2004 



239 



DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES 



which are either not current or poorly researched. It is 
not acceptable, for example for one Environmental 
Impact Statement to become the main reference for 
statements or predictions made in another such 
document, without reference to the wider and most 
current scientific literature. The following example 
from the assessment of dam development on the 
Burnett River in Queensland sharply illustrates these 
concerns. 

Arthington (2000) suggested that 
"platypusfes] feed by scooping prey items and mud 
into cheek pouches in the mouth and grinding the 
mixture to a sludge before digesting it". Based on this 
suggestion, the resultant Environmental Impact 
Statement concluded that "the deposition of sediments 
in the shallower areas of the dam would provide extra 
foraging area for Platypus [es]" as "an increase of 
available muddy substrate would provide more 
foraging area. Hard substrates offer less feeding 
opportunities because prey cannot be as easily scooped 
and ground up if they are on hard substrates or if the 
scooped material contains large pebble material" 
(Anon. 2003). Neither the literature nor the current 
study support the original suggestion by Arthington 
(2000) which consequently has led to a very equivocal 
prediction regarding the possible impact of dams on 
platypus foraging. It is crucial that such equivocal 
predictions do not become established in the literature 
consulted by those carrying out environmental impact 
assessment studies. 



ACKNOWLEDGEMENTS 

This work was carried out during monitoring 
studies associated with the Hastings District Water Supply 
Augmentation Scheme and was funded by Hastings Council 
and the New South Wales Department of Commerce Offices 
of Government Procurement and Government Business 
(formerly Department of Works and Services). Keith Bishop 
provided valuable advice during this study and comments 
on drafts of this paper. Michael Augee and another 
anonymous referee are thanked for their comments and 
recommendations on the paper. 



REFERENCES 

Anonymous. (2002). Foxes kill four platypus. Hastings 
Gazette, 19 December, 2002 p. 5. 

Anonymous. (2003) 'Burnett Catchment Water 
Infrastructure - Burnett River Dam. 
Environmental Impact Assessment. 1 1 . Aquatic 
Flora and Fauna'. (Burnett Water Pty. Ltd.: 
South Brisbane). 



Arthington, A.H. (2000). 'Burnett Basin WAMP Current 
Environmental Conditions and Impacts of 
Existing Water Resource Development'. 
Appendix H. "Reptiles, Frogs, Rats, Platypus 
and Birds' . (Queensland Department of Natural 
Resources: Brisbane). 

Bailey, N.T.J. (1969). 'Statistical Methods in Biology'. 
(The English Universities Press, London). 

Bethge, P. (2002). Energetics and foraging behaviour of 
the platypus Ornithorhynchus anatinus. PhD 
thesis. University of Tasmania, Hobart. 

Bethge, P., Munks, S. and Nicol, S. (2001). Energetics 
and locomotion in the platypus, 
Ornithorhynchus anatinus. Journal of 
Comparative Physiology - B, Biochemical, 
Systematic and Environmental Physiology 171, 
497-506. 

Bethge, P., Munks, S., Otley, H. and Nicol, S. (2003). 

Diving behaviour, dive cycles and aerobic dive 
limit in the platypus, Ornithorhynchus anatinus. 
Journal of Comparative Physiology A 136, 799- 
809. 

Boulton, A J. and Brock, M. (1999). 'Australian 
Freshwater Ecology. Processes and 
Management'. (Gleneagles Publishing: Glen 
Osmond, South Australia). 

Brooks, A.P. and Brierley, G.L. (1996). Geomorphic 

responses of the lower Bega River to catchment 
disturbance 1851-1926. Geomorphology 18, 
291-304. 

Brierley, G.J., Cohen, T., Fryirs, K, Brooks, A. 1999. 
Post-European changes to the fluvial 
geomorphology of Bega catchment, Australia: 
implications for river ecology. Freshwater 
Biology 41, 839-848. 

Bryant, A.G. (1993). An evaluation of the habitat 

characteristics of pools used by platypuses 
{Ornithorhynchus anatinus) in the upper 
Macquarie River system. Bachelor of Applied 
Science (Hons) thesis. Charles Sturt University: 
Bathurst. 

Davies, P.E. and Cook, L.S.J. (2001). 'Basslink Integrated 
Impact Assessment Statement. Potential Effects 
of changes to Hydro Power Generation' . 
Appendix 7: Gordon River Macroin vertebrate 
and Aquatic Mammal Assessment'. 

Ellem, B.A., Bryant, A. and O'Connor, A., (1998). 
Statistical modelling of platypus 
(Ornithorhynchus anatinus) habitat preferences 
using generalised linear models. Australian 
Mammalogy 20, 281-285. 

Ellem, B.A. and McLeod, A. 1998. Platypus 

(Ornithorhynchus anatinus) movement data 
from the Duckmaloi weir pool: Poisson 
regression models. Australian Mammalogy 20, 
287-292. 

Elliott, J.M. (1977). 'Some Methods for the Statistical 

Analysis of Benthic Invertebrates'. Freshwater 
Biological Association Scientific Publication 25, 
1-150. 



240 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT 



Evans, B.K. , D.R., Baldwin, J. and Gabbott, G.R.T. 1994. 
Diving ability in the platypus. Australian 
Journal of Zoology 42, 17-27. 

Faragher, R.A., Grant, T.R. and Carrick, F.N. 1979. Food 
of the platypus, Ornithorhynchus anatinus, with 
notes on the food of the brown trout, Salmo 
trutta, in the Shoalhaven River, New South 
Wales. Australian Journal of Ecology 4, 171- 
179. 

Grant, T.R. (1982). Food of the platypus, 

Ornithorhynchus anatinus (Omithorhynchidae: 
Monotremata) from various water bodies in 
New South Wales. Australian Mammalogy 5, 
135-136. 

Grant, T.R. (1993). 'The Bellinger River Water Supply 

Project Aquatic Studies - The Platypus'. (Report 
to Mitchell McCotter on behalf of the Coffs 
Harbour City Council and Department of Public 
Works by Mount King Ecological Surveys, 
Oberon). 

Grant, T.R. (1991). The biology and management of the 
platypus {Ornithorhynchus anatinus) in New 
South Wales. Species Management Report No. 
5. NSW National Parks and Wildlife Service: 
Hurstville. 

Grant, T.R. and Bishop, K.A. (1998). Instream flow 

requirements for the platypus {Ornithorhynchus 
anatinus). An assessment strategy. Australian 
Mammalogy 20, 267-280. 

Grant, T.R. and Dawson, T.J. (1978). Temperature 

regulation in the platypus, Ornithorhynchus 
anatinus, production and loss of metabolic heat 
in air and water. Physiological Zoology 51: 315- 
332. 

Grant, T.R., Griffiths, M. and Temple-Smith, P.D. (2004). 
Breeding in a Free-ranging Population of 
Platypuses, Ornithorhynchus anatinus, in the 
upper Shoalhaven River, New South Wales - a 
27 Year Study. Proceedings of the Linnean 
society of New South Wales 125, 227-236. 

Grant, T.R. and Temple-Smith, P.D. 1998. Growth of 
nestling and juvenile platypuses 
{Ornithorhynchus anatinus). Australian 
Mammalogy 20, 221-230. 



Harris, J. (1987). Growth of Australian bass, Macquaria 
novemaculeata (Perciformes; Perichthyidae), in 
the Sydney Basin, Australian Journal of Marine 
and Freshwater Research 38, 351-61 

McLeod, A. L. (1993). Movement, home range, burrow 
usage, diel activity and juvenile dispersal of 
platypus, Ornithorhynchus anatinus, on the 
Duckmaloi Weir, New South Wales' Bachelor 
of Applied Science (Hons) thesis. Charles Sturt 
University: Bathurst. 

Marchant, S. and Higgins, P.J. (eds)(1993). 'Handbook of 
Australian, New Zealand and Antarctic Birds' . 
(Oxford University Press, Melbourne). 

Olsen, P. (1999). Winged pirates. Nature Australia 26, 30- 
37. 

Rakick, R., Rakick, B., Cook, L. and Munks, S. (2001). 
Observations of a platypus foraging in the sea 
and hunting by a wedge-tailed eagle. Tasmanian 
Naturalist 123, 3-4. 

Richards, GC. 1986. predation on a platypus, 

Ornithorhynchus anatinus (Monotremata: 
Omithorhynchidae), by a goshawk. Australian 
Mammalogy 9, 67. 

Rohweder, D. (1992). Management of platypus in the 
Richmond River catchment, northern New 
South Wales. Bachelor of Applied Science 
(Hons) thesis. University of New England 
Northern Rivers: Lismore. 

Serena, M. (1994). Use of time and space by the platypus 
{Ornithorhynchus anatinus) along a Victorian 
stream. Journal of Zoology (London) 232, 117- 
131. 

Serena, M., Worley, M., Swinnerton, M. and Williams, 
G.A. (2001). Effect of food availability and 
habitat on the distribution of platypus 
{Ornithorhynchus anatinus) foraging activity. 
Australian Journal of Zoology 49, 263-277. 

Smith, A.K. and Pollard, D.A. (1998). Policy guidelines. 
Aquatic habitat management and fish 
conservation. NSW Fisheries: Sydney. 

Young, W.J. (Ed) (2001). 'Rivers as Ecological Systems: 
The Murray-Darling Basin'. (Murray-Darling 
Basin Commission: Canberra). 



Proc. Linn. Soc. N.S.W., 125, 2004 



241 



242 



Distribution of the Platypus in the Bellinger Catchment from 
Community Knowledge and Field Survey and its Relationship 

to River Disturbance 

Daniel Lunney 1 , Tom Grant 2 and Alison Matthews 1 

'Department of Environment and Conservation (NSW), PO Box 1967, Hurstville, New South Wales 2220, 
Australia; 2 School of Biological, Earth and Environmental Sciences, University of New South Wales, New 

South Wales 2052, Australia. 



Lunney, D., Grant, T. and Matthews, A. (2004). Distribution of the platypus in the Bellinger catchment 
from community knowledge and field survey and its relationship to river disturbance, Proceedings of 
the Linnean Society of New South Wales 125, 243-258. 

Platypus distribution in the Bellinger catchment was investigated using a combination of field and 
community surveys. The field survey in 1996 consisted of netting and observations from the river bank and 
a canoe. The community-based wildlife survey consisted of a questionnaire and colour maps on which 
respondents were asked to mark the locations of sightings. Platypuses were observed or caught in 36 locations 
from all three rivers of the catchment. Two of the three platypuses captured were lactating females. The 
community recorded 123 locations of platypuses. The fact that the wildlife survey yielded similar results to 
the field surveys in identifying the location of individuals highlights the value of community records for 
platypus surveys. There were major floods in 2001, after which we contacted respondents who had reported 
seeing platypuses three years before. Of the 21 respondents who had been near the river since the flood, 7 
had seen platypuses, principally in the tributaries of the Bellinger River. The habitat quality of the rivers 
was evaluated for platypuses and records were related to disturbance and rehabilitation. The species has 
survived in this system, but its future can only be assured by strategies which prevent further degradation of 
its habitat and institute proactive rehabilitation of the damaged sections of these streams. 

Manuscript received 18 August 2003, accepted for publication 8 January 2004. 

KEYWORDS: Catchment management, community wildlife survey, distribution, platypus, river 
management, wildlife. 



INTRODUCTION 

Grant (1992) and Grant et al. (2000) reported 
the distribution of the platypus Ornithorhynchus 
anatinus in New South Wales as having changed very 
little since the occupation of Australia by Europeans. 
Platypuses are considered common in the river systems 
of the coastal, tableland and western slopes (Grant 
1991, 1992) and are frequently reported from streams 
flowing through agricultural land in these areas. In 
three separate surveys in New South Wales, 52-76% 
of recorded platypus sightings were from agricultural 
land (Grant 1991; Lunney et al. 1998; Rohweder and 
Baverstock 1999). The current study investigated the 
distribution of the platypus in a typical north coast river 
system, the Bellinger catchment, where the highland 
headwater streams arise in forested areas but grade 
into predominantly agricultural land (especially cattle 
grazing) towards the coast. 

The distribution of platypuses in the 
Bellinger-Kalang river system was investigated using 



both field and community surveys. Community 
surveys have been successful in identifying locations 
of platypuses (Lunney et al. 1998; Turnbull 1998; 
Rohweder and Baverstock 1999; Otley 2001). This 
survey was part of a wider community-based survey 
of the distribution of a number of key native wildlife 
species in the Bellinger and Kalang valleys adjacent 
to Bellinger River National Park. This study sought to 
assess the co-existence of typical rural community 
activities and wildlife species, including the platypus. 
The field observations and capture of platypuses were 
compared with the questionnaire reports for this and 
other native species in the catchment with the aim of 
testing the hypothesis that information gained from 
survey data provided by the community would be a 
reliable indicator of the presence of wildlife species in 
the area. 

Since our field and questionnaire study, 
Cohen et al. (1998) assessed the Bellinger-Kalang 
catchment using the River Styles framework (Brierley 
et al. 2002) and assigned conservation and 



PLATYPUS IN THE BELLINGER CATCHMENT 



rehabilitation priorities to various stream reaches. 
Using analysis of aerial photography and field 
observations, Cohen et al. (1998) assigned various 
sections of the Bellinger-Kalang catchment to a 
number of River Styles which have particular 
geomorphic attributes. The authors of the study stress 
that these categories are a "record of the character and 
behaviour of sections of river" and "are not a direct 
measure of river condition". A separate set of 
procedures has been developed to appraise geomorphic 
river condition, building on attributes of river character 
and behaviour that are pertinent to any given River 
Style (Fryirs 2003). The attributes used in discerning 
the River Styles are shown in Table 1 and include 
channel planform and stability, morphology and 
geometry (depth and width), as well as descriptions of 
geomorphic units (e.g. pools, riffles, point bars), bed 
character (e.g. sand, gravel, cobbles) and vegetation 
character (including riparian vegetation and woody 
debris in the stream). Some of these attributes have 
been identified with the occurrence and foraging 
activity of platypuses and include: 

Channel geometry : pool depth has been 
positively related to the occurrence of the 
species (Ellem et al. 1998; Grant 2004), with 
platypuses often being observed foraging in 
water of greater than one metre depth and less 
than 5 metres. It has been suggested that 
foraging in shallow water can expose 
individuals to predation, especially from the 
introduced fox Vulpes vulpes (Grant and Denny 
1993; Serena 1994; Anon. 2002). 
Geomorphic units : pool/riffle sequences have 
also been found to be associated with the 
presence (Rohweder 1992; Bryant 1993) and 
foraging (Serena et al. 2001) of platypuses, this 
probably being related to the benthic 
productivity of such geomorphic units (Hynes 
1970; Logan and Brooker 1983; Boulton and 
Brock 1999). 

Bed character : the complexity of the bed 
substrate, including large particle sizes (rocks, 
cobbles, pebbles and gravel), has been 
positively related to both occurrence 
(Rohweder 1992) and foraging activities 
(Serena et al. 2001 ; Grant 2004) of platypuses, 
again probably resulting from greater benthic 
productivity (Hynes 1970; Marchant et al. 
1984; Boulton and Brock 1999). 
Vegetation character : medium-to-large trees, 
especially indigenous species, are associated 
with the use of river reaches by foraging 
platypuses (Serena et al. 200 1 ) and overhanging 



vegetation has also been identified as a variable 
found in areas where platypuses are found 
(Rohweder 1992; Bryant 1993; Serena et al. 
1998). This association between riparian 
vegetation and platypus occurrence is related 
to a number of important functions of such 
vegetation, including stabilisation of the bank, 
provision of cover from predators, supply of 
organic material to the food chains of the stream 
and shade moderating temperature variations, 
especially in summer (Riding and Carter 1992; 
Boulton and Brock 1999). 

The abundance of woody debris, included in 
the "vegetation character" attribute of Cohen et al. 
(1998), is also known to be positively associated with 
platypus occurrence (Rohweder 1992) and foraging 
(Serena et al. 2001), again probably being related to 
the complexity of habitats available for 
macroinvertebrates (Benke et al. 1985; Anon. 1998; 
Anon. 2000a). 

Cohen et al. (1998) also sorted sites in the 
Bellinger-Kalang catchment, grouping river reaches 
into five categories based on procedures outlined in 
Brierley and Fryirs (2000). These are summarised in 
Table 2 and are generally ranked from the least 
(conservation) to the most disturbed sites (degraded), 
although the "strategic" priority #2 sites were identified 
as being more disturbed than the priority #3 sites and 
were given a higher priority as they may impact on 
other sites downstream. This paper analyses these 
Rivers Styles and conservation/rehabilitation 
categories in relation to the data on occurrence of the 
platypus. We propose priorities for conservation and 
rehabilitation of the river system for the future survival 
of platypuses in the Bellinger and Kalang river system 
and in rural areas in general. 

Major floods in the Bellinger catchment in 
early 200 1 provided an opportunity to assess the impact 
of floods on a known population of platypuses. A 
follow-up community survey was undertaken to 
determine the survival of platypuses post-flood. 



METHODS 

Study area 

The Bellinger is a fertile river valley on the 
north coast of New South Wales just south of Coffs 
Harbour, and includes the main townships of Bellingen 
and Urunga (Figure 1). The valley extends 
approximately 50 km inland from the coast at Urunga, 
and is approximately 20 km wide from Tucker's Nob 
range in the north to the Bellbucca ridge in the south. 



244 



Proc. Linn. Soc. N.S.W., 125, 2004 



D. LUNNEY, T. GRANT AND A. MATTHEWS 



u 

2 

a 

■8 

a 
o 
•-s 

4) 
DC 
V 



"S 
s 
w 

I 

1 

© 



I 

a3 © 
it 



If 

3 3 



as 



i/i 



1 



bo ~ 

a v 

5 o 

(73 3 



•a 




o 




o 




£ 




."§ 








fC 


00 


to 


a 


8 


3 

cd 






<L> 


to 

•c 


l 


■s 


z 


T3 



co 

<3 M 

■g x> 

If 



p. . 


^ 


"O 


o 


o 


p 


a 


•a 

CD 


"2 3 

e s 


X) 
to 


"g -c 


!^ 


» co" 


ti 

CD 


a "3 
■p o 

fe a, 





2 8.8 



c « 

- "3 
co 3 

"3 is 
o .2 
p» — 



u 

43 a 

- S 

Q. TO 







4> 








T> 




4) 


p. 

CO 

*3 


1 
s 


P. 
co 

CD 

•a 


8 


S 


O 
tN 


a 


o 


"O 


i 

O 


tN 



3 



43 



(50 

a 



>. 




T3 




O 




O 




£ 




t-i 




P3 








I 




CO 




fc 










00 


to 


a 


8 

.3 


•3 

o 


I 


en 

•c 


"to 1 


X 


2 


CO 

-o 



•c 



■3 

3 g 

w . s. 



CO 



CO 

» 2 jg 

l-i *Q J5 

"'el's 

m 2 3 



CO 



tN tN 



3 
00 



3> 



co 

3 



52 

o 
"ab 

a 

t/i 



o 

* CO 

a "c 

CO 13 
P. >"» 

12 o 

S o 

8 £ 

3 2 

I §•■! 
■•a H « 

ZS5 



l-t 

o 




CO 

6 


■ a 




00 


2-g 
S2 - 
-a JJ 

•3 X> 

!■§ 


1 
60 


CO °° 
1 § 



.a -a 
s s 

P. J2 



s 



C x> 

co" "a, 

*3 *-< 

O CO 

S3 



cd 




o 




o 












T3 




CO 




CO 




m 




xj 


u 


co 


3 




Cd 


00 




a 


to 

a 


t/j 


3 



o 
O 



1-1 CO 

so 3 

.5 2 ■» 

C 3 c 

.M .3 _ 'S3 

o I -2 3. 

c o > "O 

T3 Q 5 o 

CO .co ;3 O 

CQ *3 cd 53 



co co 

o x> 

<H CO 

o-a 

5 >» 

■3 "a 

2 o 

- i 

3 5 

co a 

la" 



CO 

I- 

oo 

o "2 



1 § 

61 



.a 




o 




B, 








CO 




co 
53 


£2 


4H 


c« 


•c 


XI 






CO 


03 




a 


O 


CO 


O 




o< 


J2 




CO 




a 




CO 




60 








•a 




cd 




CO 




*-H 




Xj 


<g 




3 


— 


cd 


"ob 




.5 


to 

e 


C75 


3 



*n a "a 

^8& 



CO 


CO 


a 


■ti 


o 


X) 


« 


CO 


CO 


■a 


> 


^ 


■Jj 


•a 


ca 


o 


a 


o 


co" 


£ 


u< 


1-4 


3 


o 


to 


3 


cd 

PL, 


1 



CO CO 

cS-g 

■■a "a 
2 o 

- i 
i - 

a a 



I 

a 

00 

o 



3 



o 

•a 

05 



cd 




X) 








g 








"S 




p. 








CO 




CO 




tm 




cm 




"C 






CO 


CO 


-a 


3 


J 


o 




p, 


Cfl 



co 




T3 




'? 


P. 

co 


a 


CO 

-a 


o 

IT) 


a 


o 


en 
i 


<N 


1-C 



"3 



cd 




a 








-a 




cd 




CO 




iH 




5 


_u 


co 


3 




cd 


00 




e 
en 


to 
§ 


| 




CO 




> 




cd 




i- 




00 




00 




3 




•c 





° s 

o « 

p.- 
8 I 

rrs cd 



3 « 



Pi «S 



CO 00 

a 

cd 

X) 

2 

CO 

cd 
XI 



o 

p. 



o 
o 





CO 






•a 










o" 


'$■ 




3 a, 
2 co 

IT) c 


a 

o 
o 

T-I 






p. 


■ ■ 


o 


CO 
CO 


M tN 


en 


•a 



2 


2 




CO 


CO 




a 


. a 




CO 


co 




00 


00 










•a 


•a 




cd 


cd 




CO 


co 






t-t 




•3 


•3 


CO 


5 CO 


co 
*ob 

.a 


1 

CO 

3 


V5 to 


CZl 


3 



H 

OS 



a> 

JS 
o 



u 

ox 
a 

■ 

cu 

DC 

B 
03 



tZ3 
hi 



1 

(U 



CU 

S 



o 

S 

o 

cu 

o 



H 



■a 



Proc. Linn. Soc. N.S.W., 125, 2004 



245 



PLATYPUS IN THE BELLINGER CATCHMENT 



Priority 



Nature of Sites 



1 . Conservation sites 

2. Strategic sites 

3. High recovery potential 

4. Moderate recovery potential 

5 . Degraded 



least disturbed; river structure and vegetation relatively intact 
may be sensitive to disturbance or may affect sites downstream 
may show signs of natural recovery 
moderately degraded with reasonable potential for recovery 
highly degraded reaches with little natural recovery potential 



Table 2. Priority ranking of sites for river rehabilitation in the Bellinger- Kalang catchment (Cohen et al. 
1998). 



Forested lands rise steeply from the valley, forming 
the extremely rugged fringe of the New England 
Plateau. The Bellinger Valley comprises the catchment 
areas of the Bellinger and Kalang Rivers, referred to 
in earlier maps as the North and South Arms of the 
Bellinger River. The third major river of the valley, 
the Never Never River, is a tributary of the Bellinger 
River, which it joins near Gordonville, about 10 
kilometres upstream of Bellingen township. Tidal 
influence extends to Bellingen on the Bellinger River 
and as far as Spicketts Creek on the Kalang River 
(Cohen et al. 1998). The valley and floodplain was 



rapidly cleared by the cedar-getters and during 
settlement in the mid to late 1800s (Anon. 1978; 
Lunney and Moon 1997). Now, the land is used 
primarily for dairying and beef cattle grazing, with 
small areas being planted for crops. The population of 
the valley was 12,253 in 1996, representing a 
population growth of 21 per cent over the previous 10 
years (Anon. 2001a). 

The upper reaches of the streams of the 
Bellinger and Kalang valleys flow through steep 
forested areas in their headwaters, but degradation due 
to human activities, particularly clearing for 




Figure 1. Location map of the Bellinger catchment, showing main features of the study area and the 
sections surveyed in the Field work. 



246 



Proc. Linn. Soc. N.S.W., 125, 2004 



D. LUNNEY, T. GRANT AND A. MATTHEWS 



Location 


Kalang River 


Kalang River 


Bellinger River 




Jamisons Creek 


Jamisons Creek 


Justins Bridge 


Date 


14.12.96 


14.12.96 


15.12.96 


Sex 


Female 


Female 


Female 


Age 


adult 


adult 


adult 


Length (cm) 


39.0 


43.5 


43.0 


Bill length (cm) 


4.9 


4.9 


5.1 


Bill width (cm) 


4.2 


4.4 


4.3 


Weight (g) 


730 


905 


900 


Spur 











Milk (oxytocin) 


No 


Yes 


Yes 



Table 3. Details of platypuses captured during field survey in December 1996. 



agriculture, increases from the middle reaches to lower 
reaches upstream of the tidal limits at Bellingen and 
Spicketts Creek (Figure 1). Access of cattle to river 
banks has resulted in bank damage, especially in the 
lower Kalang River and the Bellinger River 
downstream of Thora. Parts of these sections of the 
rivers and their tributaries have good bank habitat for 
platypuses, but other sections are of lower quality due 
to the occurrence of natural gravel bars, to the effects 
of past gravel extraction, and to earth banks being 
cleared and/or damaged by cattle. The upper reaches 
of both these rivers and the Never Never River provide 
good platypus bank habitat, although some cattle 
damage to banks in parts of the upper sections of the 
Bellinger River and lower Never Never River was 
present at the time of the survey. 

Riparian vegetation is generally continuous 
on both banks of the upper reaches of all the streams 
in the system but becomes less continuous in the lower 
reaches of most streams. The Bellinger River between 
the Never Never River junction and Bellingen was 
considered to be the most degraded section of the 
system. River oak Casuarina cunninghamiana was the 
main native riparian species found, while exotic species 
- willows Salix sp., camphor laurel Cinnamomum 
camphora, privet Ligustrum sp. and lantana Lantana 
camara - were widely distributed in the riparian zones 
of most streams at the time of the study. 

Field sampling 

Field sampling was carried out over 10 days 
during December 1996. The sections of the system 
surveyed by canoe, bank observation and netting are 
shown in Figure 1 . The dates of platypus captured are 
given in Table 3. 

Canoe and bank observations were made 
either in the two hours prior to darkness and/or the 



two hours after dawn. Most of the Bellinger River from 
the mid-catchment gorge to Bellingen was surveyed 
by canoe either in the late afternoon or early morning 
(Figure 1). The Kalang River was unsuitable for canoe 
transects along much of its length due to its smaller 
size and the presence of obstacles in the channel. The 
section of river downstream from Duffys Bridge was 
suitable for use of the canoe and was surveyed a 
number of times. 

Live trapping of platypuses was carried out 
at four sites, one on the Never Never River, one on the 
upper Bellinger River and two on the Kalang River 
(Figure 1) using the methods of Grant and Carrick 
(1974). All captured females were injected with 0.2 
ml of synthetic oxytocin (Syntocinon) to indicate the 
presence of lactation (Grant and Griffiths 1992). 

The Kalang River was less intensively 
sampled than the Bellinger section of the catchment 
due to its unsuitability for canoe transects and difficulty 
of access for observation at sites which appeared to 
represent good platypus habitat. However, netting was 
carried out at one downstream and one upstream site 
on the Kalang River to assess the accuracy of reports 
obtained from local residents during the survey and to 
compare with community reports from this part of the 
river system. 

Habitat assessment 

At a number of accessible sites (mainly at 
road crossings) on the Bellinger (15), Never Never 
(8) and Kalang (15) rivers the following data or rank 
scores were collected to provide an assessment of 
habitat characteristics known to be associated with the 
occurrence of platypuses and their use of an area 
(Rohweder 1992; Bryant 1993; Ellem et al. 1998; Grant 
and Bishop 1998; Serena et al. 1998, 2001). This 
scoring procedure was based on both published and 



Proc. Linn. Soc. N.S.W., 125, 2004 



247 



PLATYPUS IN THE BELLINGER CATCHMENT 



unpublished field observations of platypus habitat: 

Habitat Category - this was a broad scoring of habitat 
suitability (1 best to 5 worst). Note: in the following 
categories, shade/shelter is usually provided by 
overhanging vegetation, but shade did not have to be 
present at time of observation as long as vegetation 
would provide shade/shelter at some times of the day. 
This is important not only to the platypus itself but to 
benthic invertebrate prey species: 

Category 1. EXCELLENT HABITAT - pools 

and/or riffle areas with >75% earth banks 

consolidated by roots of vegetation and 

providing significant shade/shelter, on both 

sides of river. 

Category 2. GOOD HABITAT - pools and/or 

riffle areas with 50-75% earth banks 

consolidated by roots of vegetation and 

providing significant shade/shelter, on at least 

one side of river or evenly distributed on both 

sides. 

Category 3. MODERATE HABITAT - pools 

and/or riffles with 25-50% earth bank 

consolidated by vegetation and providing a little 

shade/shelter. 

Category 4. POOR HABITAT - pools and/or 

riffles with 5-25% earth banks consolidated by 

roots of vegetation and providing little or no 

shade/shelter. 

Category 5. MARGINAL - pools and/or riffles 

with < 5% earth banks consolidated by roots 

of vegetation and providing no shade/shelter 



first colour map was a user-friendly map of the area 
where respondents to the survey could mark on it the 
locations of fauna, including the platypus, they had 
seen in the area. A grid was included on this map so 
that grid references could be determined with ease. 
These locations were then transferred to the 
geographical information system, ArcView, for 
analysis. The survey form, including the maps of the 
catchment, appear in Figure 2A&B. 

Relationship to River Styles 

To investigate the possibility that analysis of 
River Styles may be a useful method of predicting 
platypus occurrence or relative abundance in sections 
of a river system, platypus records from the field and 
community surveys were allocated by one of the 
authors (TRG, who has Provisional River Styler 
accreditation) to the various River Styles identified in 
the Bellinger System by Cohen et al. (1998). Stream 
reaches representing various River Styles from Cohen 
et al. (1998: Figures 1A and 9) were transposed onto 
the relevant 1:25000 topographical maps and the 
distances calculated using a manual map measure 
(Uchida Curvimeter). 

As only two platypus records were obtained 
from the mountain headwater streams and upland 
stream River Styles, these stream categories and 
observations were not included in the analysis. 
Observations of platypuses in streams which were not 
classified by Cohen et al. (1998) or were in the tidal 
sections of the rivers (one observation only) were also 
not considered. 



Riparian characteristics - these were features of banks 
that had been associated with platypus occurrence in 
other studies and were expressed as a percentage of 
sites at which they were present: 

- bank damage attributable to stock access; 

- bank damage attributable to floods; 

- presence of riparian vegetation; 

- presence of C. cunninghamiana (the most 
predominant native riparian tree species); 

- presence of introduced plant species in the 
riparian zone (especially willows, lantana, 
privet and camphor laurel). 

Community-based survey 

A community-based wildlife survey, in which 
the platypus was one of the target species, was posted 
to residents of the Bellinger-Kalang valley in 
December 1997. A total of 3000 survey forms was 
distributed by post to every household. There was a 
free-post return. The survey consisted of a 
questionnaire and colour maps on A3 size paper. The 



Relationship to river disturbance 

As was carried out for the River Styles 
categories, conservation/rehabilitation sections from 
Cohen et al. (1998: Figures 1A and 9) were transposed 
onto the relevant 1 :25000 topographical maps and the 
distances calculated using a manual map measure 
(Uchida Curvimeter). Platypus records from the field 
and community surveys were allocated to the various 
conservation/rehabilitation categories. 

Post-flood survey 

In September 2001 we contacted those 
community members who had reported platypuses in 
the wildlife survey conducted three years previously. 
A letter was individually addressed to each respondent 
and contained a covering note, a questionnaire to gather 
information on post-flood platypus sightings and a map 
showing the results of the community and field 
locations of platypuses on which each respondent could 
mark recent sightings. 



248 



Proc. Linn. Soc. N.S.W., 125, 2004 



D. LUNNEY, T. GRANT AND A. MATTHEWS 



Bellinger Valley 
Wildlife Survey 




No Postage stamp required 
if posted in Australia 




Please fold and return to: 



Reply Paid 100 

Bellinger Valley Wildlife Survey 

c\- Dan Lunney 

Biodiversity Survey and Research Division 

NSW National Parks and Wildlife Service 

PO Box 1967 

HURSTVILLE NSW 2220 




Dear Shire Resident or Visitor, 

We are seeking your co-operation in conducting a wildlife survey of the Bellinger 
Valley. Its purpose is to locate wildlife populations as well as the habitats that 
are important for them. The long-term aim is to improve wildlife management of 
the valley by knowing which animals inhabit the area, where they occur, and the 
possible threats to their survival. This survey has the endorsement of Bellingen 
Shire Council and is supported by grants from the Heritage Assistance Program 
and the Foundation for National Parks and Wildlife. 

We would like you to fill out this survey even if you have only one wildlife 
sighting to record or you can only complete a part of the form. Also, if you have 
any historical information, this would help us understand the changes that have 
occurred to local wildlife populations over time in the Bellinger Valley. 

Please post your completed survey form (no stamp required) by 16 February 
1998. 

Thank you for taking the time to assist us in compiling this community-based 
survey. If you would like a souvenir copy of this form, please tick the box on 
page 4. 




Dan Lunney 

(02) 9585 6489 



Alison Matthews 

(02) 9585 6559 



Dionne Coburn 

(02) 9585 6558 



New South Wales National Parks and Wildlife Service 
December 1997 



NSW 

NATIONAL 
PARKS AND 
WILDLIFE 
SERVICE 



Figure 2A. The Bellinger Wildlife Survey form. 



Proc. Linn. Soc. N.S.W., 125, 2004 



249 



PLATYPUS IN THE BELLINGER CATCHMENT 




Figure 2B. The Bellinger catchment map provided with the Wildlife Survey form. 



250 



Proc. Linn. Soc. N.S.W., 125, 2004 



D. LUNNEY, T. GRANT AND A. MATTHEWS 



RESULTS 

Field survey 

Platypuses were observed or caught in all 
three rivers at a total of 36 locations (Figure 3). Two 
platypuses were captured at one of the two sites (the 
upstream site) on the Kalang River but none was 
observed in the limited sampling of this river by foot 
or by canoe. None was caught at the Never Never River 
netting site, but one was captured at the site on the 
upper Bellinger River. All the individuals captured 
were female, two of which were lactating, indicating 
the occurrence of breeding populations in both the 
Bellinger and Kalang rivers. All animals captured were 
within expected dimensions and body condition (Table 
3). Platypuses were found to be common and 
continuously distributed along the Bellinger and Never 
Never Rivers, being captured or observed at 35 sites 
(Figure 3). 

The canoe transect survey method was most 
successful, yielding 2.2 animals per hour of 
observation, compared with 0.17 for both netting and 



observations by foot from river banks (Table 4). 

Community reporting of the occurrence of 
platypuses 

A total of 522 replies (17.4% return) was 
received to the Bellinger valley wildlife survey. 
Platypuses were recorded at 123 sites by the 
community-based survey. Only two platypus records 
were obtained from the headwater streams of the 
catchment and the field survey did not sample these 
streams. These data showed a much lower number of 
sightings (13) in the Kalang River and its tributaries 
than in the Bellinger River (110) and its tributary 
streams. 

The field and community-based data showed 
that the platypus is commonly found throughout the 
Bellinger River catchment, including the Never Never 
and Rosewood Rivers, and its distribution is probably 
continuous above the tidal limit at Bellingen to the 
headwater streams, which were not surveyed in this 
study. There was one report of a platypus downstream 
of the tidal limit on the Bellinger River. As well as 
being reported less often along the Kalang River 



A/ River 
'[] Bellingen LGA 
| 3] National Parks 
| ITT1 State Forests 

* Platypus field records 

* Platypus community records 
All community records 




Figure 3. The location of field and community-based records of platypuses in the Bellinger catchment. 



Proc. Linn. Soc. N.S.W., 125, 2004 



251 



PLATYPUS IN THE BELLINGER CATCHMENT 



Method 


Kalang River 




Bellinger River 


Never Never River 


Total 




Hrs No. 


CPU 

* 


Hrs No. CPU 


Hrs 


No. 


CPU 


CPU 


Canoe 


3.75 





95 29 3.05 


2.75 


6 


2.2 


2.20 


observations 
















Bank 


4.0 





0.25 1 4.0 


15 








0.17 


observations 
















Netting 


8.5 2 


0.23 


45 1 0.22 


5 








0.17 


Total 


16.25 2 


0.12 


14.25 31 2.18 


9.25 


6 


0.65 


0.98 



*CPU: Catch/Observation per Unit Effort 

Canoe = individuals seen/hour observation in each observation period 

Observation = individuals seen/hour in each observation period 

Netting = individuals captured per net hour (1x50m net in water for 1 hour) 

Total = division of total individuals seen/caught by total hours of observation or net hours 

Table 4. Success of various field survey methods used for recording platypuses in December 1996. 



system, platypuses seemed to be more discontinuous 
in their distribution in this part of the river system 
(Figure 3). 

Reliability of the community-based data set 

The distribution of community-based reports 
of platypuses in the Kalang and Bellinger components 
of the river system showed considerable overlap with 
the field records (Figure 3). There were few 
observations by the community outside the areas in 
which the field work identified the occurrence of the 



species. One exception to this was the section of the 
Kalang River between Moodys Bridge and Sunny 
Corner, where no sightings or captures were made 
during the field work, but where platypuses were 
reported by the community. 

Habitat assessment 

There was little difference among the 15 sites 
sampled on the Bellinger and Kalang rivers in terms 
of habitat suitability (Figure 4), although on the Kalang 
River, 13% of the sites sampled were classified as 



90 
80 
70 - 



4- 

o 


60 


DC 




<u 




at 


bO 


U 




u 


40 


Ol 




o- 




* 


30 




20 




10 








1 




■ Kalang 
□ Bellinger 

M Never Never 



2 3 4 

Bank Categories 



Figure 4. Percentage of sites at which bank suitability categories were recorded on the Kalang, Bellinger 
and Never Never Rivers. 



252 



Proc. Linn. Soc. N.S.W., 125, 2004 



D. LUNNEY, T. GRANT AND A. MATTHEWS 



120 n 




■ Kalang 
□ Bellinger 

M Never Never 



Cattle Flood Riparian Casuarinas Introduced 

Damage Damage Vegetation. Plants 

Bank Characteristics 



Figure 5. Percentage of sites exhibiting various bank characteristics on the Kalang, Bellinger and Never 
Never Rivers. 



category 1, whereas none of the sites on either the 
Bellinger River or its major tributary, the Never Never 
River, fell into this category. 

All sites sampled had some riparian 
vegetation present but fewer sites on the Bellinger 
River had introduced species of riparian plants and 
more had C. cunninghamiana trees on the riverbank. 
More sites on the Kalang and Never Never Rivers than 
on the Bellinger River exhibited cattle and flood 
damage (Figure 5). 

Relationship to River Styles 

Table 5 details the lengths of each River Style, 
the total numbers of platypuses recorded in the field 
and community surveys and the numbers of platypus 
records per kilometre of each River Style. 

The River Styles of the Bellinger and Kalang 
sections of the catchment differ, with almost all (97%) 
of the Kalang River catchment being classified as 
confined bedrock with discontinuous alluvial 
floodplains, while the Bellinger River catchment 
contained a variety of Rivers Styles, ranging from 57% 
confined bedrock with discontinuous alluvial flood 
plains, through 24% alluvial with a meandering gravel 
bed to 10% and 9% of alluvial stream with a wandering 
gravel or discontinuous bed (Table 5). 

In the Bellinger River, there were 



significantly more platypus records in the alluvial 
meandering gravel bed sections of river and fewer in 
the discontinuous alluvial stream than expected if 
platypuses were distributed uniformly across River 
Styles (% 2 =26.64, 3d.f., P<0.01). In the Kalang River, 
platypus records were distributed evenly across River 
Styles. Expressed on the basis of platypus records per 
kilometre of river represented by each River Style, the 
Bellinger River had 0.95 records/km in the confined 
bedrock with discontinuous flood plains River Style, 
while the Kalang River (where this River Style made 
up 97% of the river downstream of the mountainous 
headwater reaches) had only 0.23 records/km (less than 
25% of the value for the Bellinger). In the only other 
River Style represented in the Kalang River, alluvial 
river with meandering gravel bed (3% of the river), 
there were no platypus records and yet this was the 
River Style on the Bellinger River which had most 
records (2.1/km). 

Relationship to river disturbance 

The lengths of each conservation and 
rehabilitation priority reaches proposed by Cohen et 
al. (1998), along with the total numbers of platypus 
records from the field and community-based data, as 
well as the number of records per kilometre for each 
priority category in the Bellinger and Kalang 



Proc. Linn. Soc. N.S.W., 125, 2004 



253 



PLATYPUS IN THE BELLINGER CATCHMENT 







Bellinger River 




Kalang River 




River Style 


Distance 


Platypus Platypus 


Distance 


Platypus 


Platypus 




(km) 


per km 


(km) 




per km 


Confined bedrock 


58.9 


56 0.95 


59.9 


14 


0.23 


with discontinuous 












floodplain 












Alluvial meandering 


24.8 


53 2.14 


2.1 








gravel bed river 












Alluvial wandering 


9.8 


13 1.33 











gravel bed 












Discontinuous 


9.1 


3 0.33 











alluvial stream 












Total 


102.6 


125 122 


62 


14 


023 



Table 5. River Style distances, numbers of platypus records and numbers of platypuses reported per 
kilometre of river in the Bellinger and Kalang rivers and their tributaries in December 1996. 



catchments, were compared (Table 6). The number of 
platypus records per kilometre of river in the Bellinger 
River increased from the "strategic" category (0.74/ 
km) through the "high recovery potential" (0.97/km) 
and "moderate recovery potential" (1.80/km) 
categories to be highest in the most "degraded" (2.25/ 
km) section of the river. There were significantly more 
platypus records than expected in the degraded and 
moderate recovery potential categories (x 2 =17.99, 
3d.f, p<0.01). In addition, the number of records of 
platypuses in the Bellinger River was much higher 
(1.22/km) than in the Kalang River (0.23/km), in spite 
of the fact that the latter system appears to be less 
disturbed than the Bellinger River. 



Post-flood survey 

A total of 43 replies was received from 
respondents who had reported platypuses in the 1997 
survey. Twenty-one respondents had been near the 
river since the 2001 floods and of these, 7 had seen 
platypuses. Sightings of platypuses post-flood were 
in Hydes Creek (6 sightings), Boggy Creek (1 
sighting), the Never Never River (1 sighting), the 
Kalang River between Duffys and Moody s bridges (1 
sighting) and the upper Bellinger River between 
Diehappy and Bishops Creeks (2 sightings). 





Bellinger River 




Kalang River 




River Style 


Distance Platypus 


Platypus 


Distance 


Platypus 


Platypus 




(km) 


per km 


(km) 




per km 


Conservation 


- 


- 


20.5 


5 


0.24 


Strategic 


5.4 4 


0.74 


13.9 


4 


0.29 


High recovery 


70.9 69 


0.97 


21.9 


5 


0.22 


potential 












Moderate recovery 


16.1 29 


1.80 


5.7 








potential 












Degraded 


10.2 23 


2.25 


- 


- 


- 


Total 


102.6 125 


122 


62 


14 


023 



Table 6. Conservation/rehabilitation priority distances, numbers of platypus records and numbers of 
platypuses reported per kilometre of river in the Bellinger and Kalang rivers and their tributaries in 
December 1996. 



254 



Proc. Linn. Soc. N.S.W., 125, 2004 



D. LUNNEY, T. GRANT AND A. MATTHEWS 



DISCUSSION 

Data from the field and community surveys 

The results strongly support the hypothesis 
that the community-based data from this study are 
reliable. The field observations and captures of 
platypuses closely corresponded to community 
records. As a result of this correspondence, the field 
and community data were combined in the analysis of 
platypus occurrence in relation to River Styles and 
conservation and rehabilitation priorities. Further, a 
post-flood survey was able to be conducted because 
of the reliability of community records. 

The lack of sightings from the headwater 
streams of the whole catchment suggest a lack of 
observation, rather than the species not occurring in 
them. This is supported by the low reporting of other 
wildlife species in the community wildlife survey in 
these areas (Figure 3). It is assumed that platypuses 
would almost certainly occupy these sections of the 
catchment streams, although this was not determined 
by the field survey. 

The field and community-based data rank the 
Bellinger River part of the system as being more 
suitable for occupation by platypuses than the Never 
Never and Kalang rivers. The limited habitat data 
collected during the survey point to the Kalang River 
having less suitable platypus habitat than either the 
Never Never or Bellinger Rivers. The discontinuous 
distribution of the platypus in the Kalang River, 
especially between Moodys Bridge and Rosewood 
Creek, almost certainly identified poorer habitat 
conditions. As community reports of other wildlife 
species were made along the Kalang River, the lack of 
platypus records from these sections means that the 
species is not found or is uncommon in these sections 
of the river. Further, any temporary loss of individuals 
from an area, such as from a flood, should not affect 
an accurate determination of distribution if the 
community had observed them in these sections of the 
river at other times. This is one of the values of the 
community survey, namely it was not restricted to one 
point in time and it also considered historical records. 
Qualitative field observations of the Kalang River 
between Moodys Bridge and Rosewood Creek 
confirmed that this section had poorer habitat quality 
than other sections of the river, with considerable 
disturbance of the river banks due to depletion of the 
riparian vegetation and cattle access, as well as 
accumulation of sand in the river bed. 

The field survey did not allow an adequate 
explanation to be made of the differences between the 
Bellinger and Kalang sections of the river system in 
terms of the observed platypus distribution. While parts 



of the Kalang were more degraded than sections of 
the Bellinger and Never Never rivers, platypus reports 
and field sightings were common in the Bellinger River 
between the Never Never River junction and the town 
of Bellingen, a river section which was also highly 
altered by bank clearing, stock damage to banks and 
past gravel extraction. 

One report was obtained of a platypus in the 
tidal section of the Bellinger River, close to the entrance 
of Connells Creek. Three reports of platypuses in the 
tidal section of the river were also made to the authors 
during the field study; one at Fernmount in the 1940s, 
another at the mouth of Hydes Creek in 1996 and one 
near the Old Butter Factory in Bellingen (which was 
said to be "recent"). Platypuses have occasionally been 
found in the sea (Fleay 1980; Connolly and Obendorf 
1998) and in estuarine habitats, but such occurrences 
are irregularly reported and are considered unusual 
(Stone 1983; Grant 1991, 1999; Rohweder 1992; Hird 
1993; Menkhorst 1995; Connolly and Obendorf 1998; 
Rakick et al. 2001). It seems unlikely that the species 
regularly occupies the brackish or saline waters of 
estuarine environments. Nothing is known of its 
abilities to osmoregulate under marine or brackish 
conditions or any need by the species to have access 
to fresh water to groom salt from the fur, as occurs in 
several species of otters (Kruuk 1995). Platypuses are 
known to consume a range of benthic invertebrates as 
food but insect larvae are the most common prey items 
(Faragher et al. 1979; Grant 1982). In a number of 
rivers along the coast of New South Wales, tidal 
influences and/or saline intrusion into the lower reaches 
results in the diversity of benthic macroinvertebrates 
beginning to change at the tidal limit from being 
numerically dominated by insect fauna to being 
dominated by Crustacea, including amphipods and 
isopods, with oligochaetes worms and gastropod 
molluscs also having greater representation 
(Anon. 1993 ; Simon Williams, then of Australian Water 
Technologies, pers. comm.). This could affect platypus 
distribution in the lower reaches of rivers of coastal 
New South Wales. It is also known that increased 
conductivity impairs the ability of the platypus to locate 
moving prey items, particularly small invertebrates, 
using the electrosensory mechanisms in its bill 
(Pettigrew et al. 1998). Competition with benthic- 
feeding fish species, which do not enter the freshwater 
sections of rivers, and possible predation by larger fish 
species, may also be involved in the occurrence of 
platypuses being unusual in tidal areas. 

Relationship to River Styles 

The differences in distribution and numbers 
of platypus records between the two rivers were not 



Proc. Linn. Soc. N.S.W., 125, 2004 



255 



PLATYPUS IN THE BELLINGER CATCHMENT 



found to be related to the differences in River Styles 
between the two parts of the system. On the basis of 
our findings in the Bellinger catchment we consider 
that analyses using the River Styles framework 
(Brierley et al. 2002) will not successfully predict the 
occurrence of platypuses. However, methods 
integrating geomorphic and biological considerations 
could lead to a framework which may be capable of 
predicting the suitability of streams for occupation by 
the platypus. Such integration could also provide a 
better basis for river management and rehabilitation 
than arises from the consideration of either geomorphic 
or biological considerations in isolation. This approach 
has been called the "landscape ecology approach" by 
Tockner et al. (2002). 

Relationship to river disturbance 

This study has shown platypuses to be present 
in degraded habitat of the Bellinger catchment. 
However, it would be a mistake to be complacent about 
these observations and regard disturbances of rivers 
to be benign with respect to the platypus. 

Despite the common occurrence of platypuses 
in agricultural areas, there are strong indications that 
platypus distribution has been fragmented and/or their 
numbers reduced in the streams of the Eden area 
(Lunney et al. 1998) and in the Bega (Brooks and 
Brierley 1997), Thredbo (Goldney 1998) and 
Richmond (Rohweder and Baverstock 1999) rivers of 
New South Wales and in the Wimmera River system 
in Victoria (Anon. 1999, 2000b, 2001b). In each of 
these instances the changes have been mainly attributed 
to the effects of agricultural practices. Lack of reports 
of platypuses from the Kalang River in the disturbed 
section between Moodys Bridge and Rosewood Creek 
also point to a fragmentation of platypus distribution 
within this part of the Bellinger catchment. 

Lunney et al. (1998) attributed fragmentation 
of platypus populations in the Eden region (Bega 
Valley Shire) of New South Wales to the effects of 
farming, particularly cattle grazing and the clearing 
of the riparian vegetation since 1830. Brooks and 
Brierley (1997) and Brierley et al. (1999) have detailed 
the effects of early agricultural practices in the Bega 
River valley of New South Wales, confirming that 
these practices were almost certainly responsible for 
the irreversible changes to that river system. However, 
Turnbull ( 1 998) recorded the occurrence of platypuses 
in most of the rivers around Bombala, in the tableland 
headwater streams of the Bega and Snowy Rivers in 



New South Wales, in spite of the area having been 
utilised for both cattle and sheep grazing for the past 
160 years. 

Of the 11 platypus sightings made by 
respondents to the post-flood questionnaire in the 
Bellinger catchment, 8 were in tributary streams. This 
suggests that the tributaries act as refuge areas during 
extreme floods. The tributaries could also be important 
for this population if the main streams of the Bellinger 
River system experience further degradation. This 
latter suggestion is based on our data from the Bega 
River (Lunney et al. 1998) where historically 
platypuses were found in the lower reaches, but it is 
now so degraded, shallow, sandy and exposed, that it 
no longer supports viable platypus populations. Instead 
platypuses occur only in the more protected and less 
developed tributary streams of the Bega River system. 

Conservation and rehabilitation 

Considering that the distribution of this 
unique Australian species overlaps extensively with 
activities of rural communities, its conservation 
depends on the adaptive management of these 
activities. The species has survived the current 
environmental disturbances so far, but its future 
conservation can only be assured by strategies aimed 
at preventing any further degradation of its habitat in 
these areas and by proactive rehabilitation of damaged 
sections of streams and a recognition of the possible 
importance of the tributary streams in retaining refuge 
populations of platypuses. 



ACKNOWLEDGEMENTS 

The authors are indebted to many people, 
particularly those who took the time to respond to the survey. 
We wish to acknowledge the contribution of P. Sherratt for 
the design of the community survey form; K. Weinman for 
production of the map and extra digitising of road and river 
systems that were not previously available on the National 
Parks and Wildlife Service geographic information system; 
and I. Dunn and G. Brierley for critical comments on the 
manuscript. Funding was provided by the Foundation for 
National Parks and Wildlife and a National Estate Grant. 
Platypuses were captured under licences issued by the NSW 
National Parks and Wildlife Service (A 184) and NSW 
Fisheries (Scientific Research Permit F84/1245) and under 
Ethics Approval from the University of NSW Animal Care 
and Ethics Committee (Animal Research Authority ACE 94/ 
91). 



256 



Proc. Linn. Soc. N.S.W., 125, 2004 



D. LUNNEY, T. GRANT AND A. MATTHEWS 



REFERENCES 

Anonymous. (1978). Pioneering in the Bellinger Valley. 

(The Bellinger Courier-Sun, Bellingen). 
Anonymous. (1993). 'Investigation of the invertebrate 

benthic fauna of the Bellinger River' . 

(Unpublished report for Mitchell McCotter and 

Associates. June, 1993). 
Anonymous. (1998). 'Riparian management #7. Managing 

snags in rivers'. (Land and Water Resources 

Research and Development Corporation, 

Canberra). 
Anonymous. (1999). Wimmera update. Newsletter of the 

Australian Platypus Conservancy Ripples 13, 2. 
Anonymous. (2000a). 'Snags. A valuable but scarce 

resource'. (CRC for Freshwater Ecology, 

Canberra). 
Anonymous. (2000b). Barwon platypus study. Newsletter 

of the Australian Platypus Conservancy Ripples 

16, 2. 
Anonymous (2001a). 2000-2001 State of Environment 

Report Bellingen LGA. Bellingen Shire 

Council. 
Anonymous. (2001b). Recent APC platypus survey 

results: Hopkins and Wimmera rivers. 

Newsletter of the Australian Platypus 

Conservancy Ripples 18, 2. 
Anonymous (2002). 'Foxes kill four platypus'. Hastings 

Gazette, 19 December 2002, p. 5. 
Benke, A.C., Henry, R.L., Gillespie, D.M. and Hunter, 

R.J. (1985). Importance of snag habitat for 

animal production in south eastern streams. 

Fisheries 10, 8-13. 
Boulton, A.J. and Brock, M.A. (1999). 'Australian 

Freshwater Ecology. Processes and Management' . 
(Gleneagles Publishing, Glen Osmond, South 
Australia). 
Brierley, G.J., Cohen, T., Fryirs, K. and Brooks, A. 

(1999). Post-European changes to the fluvial 

geomorphology of Bega catchment, Australia: 

implications for river ecology. Freshwater 

Biology 41, 839-848. 
Brierley, G.J. and Fryirs, K. (2000). River Styles, a 

geomorphic approach to catchment 

characterisation: implications for river 

rehabilitation in the Bega catchment, New South 

Wales, Australia. Environmental Management 

25, 661-679. 
Brierley, G., Fryirs, K., Outhet, D. and Massey, C. (2002). 

Application of the River Styles framework as a 

basis for river management in New South 

Wales, Australia. Applied Geography 22, 91- 

122. 
Brooks, A. P. and Brierley, G.J. (1997). Geomorphic 

responses of lower Bega River to catchment 

disturbance, 1851-1926. Geomorphology 18, 

291-304. 
Bryant, A.G. (1993). 'An evaluation of the habitat 

characteristics of pools used by platypuses 

(Ornithorhynchus anatinus) in the upper 



Macquarie River system'. (Bachelor of Applied 
Science (Hons) Thesis, Charles Sturt University, 
Bathurst). 

Cohen, T., Reinfelds, I. and Brierley, G. (1998). 'River 
styles in the Bellinger-Kalang catchment'. 
(Report completed for NSW Department of 
Land and Water Conservation. September, 
1998. School of Earth Sciences, Macquarie 
University, Macquarie Research Limited). 

Connolly, J.H. and Obendorf, D.L. (1998). Distribution, 
captures and physical characteristics of the 
platypus {Ornithorhynchus anatinus) in 
Tasmania. Australian Mammalogy 20, 231-237. 

Ellem, B.A., Bryant, A. and O'Connor, A. (1998). 
Statistical modelling of platypus 
(Ornithorhynchus anatinus) habitat preferences 
using generalised linear models. Australian 
Mammalogy 20, 281-285. 

Faragher, R.A., Grant, T.R. and Carrick, F.N. (1979). 
Food of the platypus, Ornithorhynchus 
anatinus, with notes on the food of the brown 
trout, Salmo trutta, in the Shoalhaven River, 
New South Wales. Australian Journal of 
Ecology 4, 171-179. 

Fleay, D. (1980). 'Paradoxical platypus. Hobnobbing with 
duckbills'. (Jacaranda Press, Milton, 
Queensland). 

Fryirs, K. (2003). Guiding principles for assessing 

geomorphic river condition: application of a 
framework in Bega catchment, South Coast, 
New South Wales, Australia. Catena 53, 17-52. 

Goldney, D. (1998). The distribution and abundance of 

platypuses in the Thredbo River-Lake Jindabyne 
system. Australian Mammalogy 20, 302-303. 

Grant, T.R. (1982). Food of the platypus, 

Ornithorhynchus anatinus (Ornithorhynchidae: 
Monotremata) from various water bodies in 
New South Wales. Australian Mammalogy 5, 
135-136. 

Grant, T.R. (1991). 'The biology and management of the 
platypus {Ornithorhynchus anatinus) in New 
South Wales'. Species Management Report No. 
5. (NSW National Parks and Wildlife Service, 
Hurstville). 

Grant, T.R. (1992). Historical and current distribution of 
the platypus, Ornithorhynchus anatinus, in 
Australia. In 'Platypus and echidnas' (Ed. M.L. 
Augee). pp. 232-254. (Royal Zoological Society 
of NSW, Sydney). 

Grant, T.R. (1999). Distribution of platypuses and 

platypus habitat in the Richmond River estuary 
and investigation of salinity tolerance in the 
species. In 'An investigation of the potential 
ecological impacts of freshwater extraction from 
the Richmond River tidal pool' (Eds. W.L. 
Peirson, K.A Bishop, R. Nittim and M.J. 
Chadwick) Appendix C. (Water Research 
Laboratory Technical Report 99/51, University 
of New South Wales, Sydney). 



Proc. Linn. Soc. N.S.W., 125, 2004 



257 



PLATYPUS IN THE BELLINGER CATCHMENT 



Grant, T.R. (2004). Depth and substrate selection by 

platypuses, Omithorhynchus anatinus, in the 
Lower Hastings River, New South Wales. 
Proceedings of the Linnean Society of New 
South Wales (submitted for this volume). 

Grant, T.R. and Bishop, K.A. (1998). Instream flow 

requirements for the platypus (Omithorhynchus 
anatinus). An assessment strategy. Australian 
Mammalogy 20, 267-280. 

Grant, T.R. and Carrick, F.N. (1974). Capture and 
marking of the platypus, Omithorhynchus 
anatinus, in the wild. Australian Zoologist 18, 
133-135. 

Grant^ T.R. and Denny, M.J.S. (1993). 'The Bellinger 

river water supply project - Aquatic studies - the 
platypus'. Raising of Karangi Dam, 
Environmental Impact Statement, 
Supplementary Documents. (Mitchell McCotter, 
Sydney). 

Grant, T.R., Gehrke, P.C., Harris, J.H. and Hartley, S. 
(2000). Distribution of the platypus 
(Omithorhynchus anatinus) in New South 
Wales: Results of the 1994-96 New South 
Wales Rivers survey. Australian Mammalogy 
21, 177-184. 

Grant, T.R. and Griffiths, M. (1992). Aspects of lactation 
and determination of sex ratios and longevity in 
a free-ranging population of platypuses, 
Omithorhynchus anatinus, in the Shoalhaven 
River, New South Wales. In 'Platypus and 
echidnas' (Ed. ML. Augee) pp. 80-89. (Royal 
Zoological Society of NSW, Sydney). 

Hird, D. (1993). Estuarine platypus activity. Tasmanian 
Naturalist 144, 7-8. 

Hynes, H.B.N. (1970). 'The ecology of running waters'. 
(University of Toronto Press, Toronto). 

Kruuk, H. (1995). 'Wild otters, predation and 

populations'. (Oxford University Press, 
Oxford). 

Logan, P. and Brooker, M.P. (1983). The 

macroinvertebrate faunas of riffles and pools. 
Water Research 17, 263-270. 

Lunney, D. and Moon, C. (1997). Flying foxes and their 
camps in the remnant rainforests of northeastern 
New South Wales. In 'Australia's Ever Changing 
Forests III' (Ed. J. Dargavel) pp. 247-277. (CRES, 
Canberra). 

Lunney, D., Grant, T., Matthews, A., Esson, C, Moon, C. 
and Ellis, M. (1998). Determining the 
distribution of the platypus (Omithorhynchus 
anatinus) in the Eden region of south-eastern 
New South Wales through community-based 
surveys. Australian Mammalogy 20, 239-250. 

Marchant, R. Mitchell, P. and Norris, R. (1984). A 

distribution of benthic invertebrates along a 
disturbed section of the La Trobe River, 
Victoria: An analysis based on numerical 
classification. Australian Journal of Marine and 
Freshwater Research 35, 355-374. 



Menkhorst, P.W. (ed.) (1995). 'Mammals of Victoria. 
Distribution, ecology and conservation'. 
(Oxford University Press, Melbourne). 

Otley, H.M. (2001). The use of a community-based survey 
to determine the distribution of the Platypus 
Omithorhynchus anatinus in the Huon River 
catchment, southern Tasmania. Australian 
Zoologist 31, 632-641. 

Pettigrew, J.D., Manger, P.R. and Fine, S.L.B. (1998). 
The sensory world of the platypus. 
Philosophical Transactions of the Royal Society 
of London B. 353, 1199-1210. 

Rakick, R., Rakick, B., Cook, L. and Munks, S. (2001). 
Observations of a platypus foraging in the sea 
and hunting by a wedge-tailed eagle. Tasmanian 
Naturalist 123, 3-4. 

Riding, T. and Carter, R. (1992). 'The importance of the 
riparian zone in water resource management - a 
literature review'. (Department of Water 
Resources, Sydney). 

Rohweder, D. (1992). 'Management of platypus in the 
Richmond River catchment, northern New 
South Wales'. (Bachelor of Applied Science 
(Hons) Thesis, University of New England 
Northern Rivers, Lismore). 

Rohweder, DA. and Baverstock, P.R. (1999). Distribution 
of platypus, Omithorhynchus anatinus, in the 
Richmond River Catchment, northern New 
South Wales. Australian Zoologist 31, 30-37. 

Serena, M. (1994). Use of time and space by platypus 
(Omithorhynchus anatinus,: Monotremata) 
along a Victorian stream. Journal of Zoology 
(London) 232, 117-131. 

Serena, M., Thomas, J.L., Williams, GA. and Officer, 

R.C.E. (1998). Use of stream and river habitats 
by the platypus, Omithorhynchus anatinus, in 
an urban fringe habitat. Australian Journal of 
Zoology 46, 267-282. 

Serena, M., Worley, M., Swinnerton, M. and Williams, 
GA. (2001). Effect of food availability and 
habitat on the distribution of platypus 
(Omithorhynchus anatinus) foraging activity. 
Australian Journal of Zoology 49, 263-277. 

Stone, G.C. (1983). 'Distribution of the platypus, 

Omithorhynchus anatinus, in Queensland'. 
(Queensland National Parks and Wildlife 
Service, Brisbane). 

Tockner, K. Ward, I.V., Edwards, P.J. and Kollman, J. 
(2002). Riverine landscapes: an introduction. 
Freshwater Biology 47, 497-501. 

Turnbull, R.W. (1998). Distribution of the platypus 

(Omithorhynchus anatinus) in the Bombala 
River catchment, South-Eastern New South 
Wales. Australian Mammalogy 20, 251-256. 



258 



Proc. Linn. Soc. N.S.W., 125, 2004 



Reducing the By-catch of Platypuses {Ornithorhynchus 
anatinus) in Commercial and Recreational Fishing Gear in New 

South Wales 

T.R. Grant 1 , M.B. Lowry 2 , Bruce Pease 2 , T.R. Walford 2 and K. Graham 2 

1 School of Biological, Earth and Environmental Sciences, University of New South Wales, Kensington NSW 
2052. email: t.grant@unsw.edu.au 2 NSW Fisheries P.O. Box 21 Cronulla, NSW 2230. 

Grant, T.R., Lowry, M.B., Pease, B., Walford, T.R. and Graham, K. (2004). Reducing the by-catch of 
platypuses {Ornithorhynchus anatinus) in commercial and recreational fishing gear in New South Wales. 
Proceedings of the Linnean Society of New South Wales 125, 259-272. 

The problem of platypus by-catch mortality in the eel, yabby and carp trap fisheries in New South 
Wales is reviewed, and the results of several experiments to determine the effectiveness of gear modifications 
to reduce platypus by-catch are presented. Entrance screens with 50-60 mm openings prevented the entry of 
platypuses into eel or yabby traps. Larger screens were not effective as a deterrent to platypuses entering 
traps. By-catch of platypuses in the eel fishery can be minimised by restricting traps to estuarine areas, 
where platypuses seldom occur, and by providing air spaces in the cod ends of traps used in impoundments 
and farm dams. Prohibiting the use of yabby traps in areas where platypuses are known to occur provides 
the most practical protection against by-catch of platypuses in this fishery. Platypuses were unable to exit 
from prototype carp traps, designed to permit escape of air-breathing species, but the provision of 
appropriately-sized openings at the base of the entrance funnels in these drum traps permitted platypuses to 
escape. 

Manuscript received 4 September 2003, accepted for publication 24 November 2003. 

KEYWORDS: by-catch, carp, eel, fishing, Ornithorhynchus anatinus, platypus, yabby. 



INTRODUCTION 

By-catch mortality of air-breathing 
vertebrates, including several species of freshwater 
turtles and diving birds, water rats (Hydromys 
chrysogaster) and platypuses {Ornithorhynchus 
anatinus), has been recognised for some time as a 
significant problem in various inland fisheries in 
Australia (Jackson 1979; Beumer et al. 1981; Grant 
1991, 1993; Grant and Denny 1991; Leadbitter 2001). 
Such by-catch mortality of platypuses is of particular 
concern in small streams, where multiple drownings 
of breeding individuals have the potential to impact 
severely on small local populations. For example, an 
abandoned fyke net in a tributary of the Gellibrand 
River in Victoria was found to contain the skeletons 
of 17 platypuses (Serena 2003). 

There has often been conflict between the 
desires of fishers to maximise catches of their target 
species, and the implementation of effective methods 
to reduce non-target by-catch. This has resulted in a 
diverse range of regulations enacted by fishery 
authorities and voluntary gear modifications by fishers 
aimed at reducing the mortality of non-target species 
(e.g. Leadbitter 2001). Unfortunately, little research 



or monitoring has been done to assess the effectiveness 
of voluntary and regulated gear modifications. 

An historical assessment of inland fishing in 
New South Wales showed that commercial fishing 
probably resulted in significant platypus mortality 
when small-mesh nets were used (Grant 1991, 1993; 
Grant and Denny 1 99 1 ). No commercial or recreational 
fishery using nets or traps to capture native fish species 
or salmonids in freshwater sections of coastal rivers is 
now permitted in New South Wales (NSW), but there 
is a commercial eel fishery based on the use of baited 
traps in estuaries, farm dams and a few large 
impoundments. West of the Great Dividing Range, the 
commercial fishery for native fin-fish species was 
phased out in 2001 . Fishers previously involved in that 
industry have been encouraged to fish for yabbies, 
mainly (Cherax destructor), using "Opera house" traps 
(Rankin 2000). The introduced carp (Cyprinus carpio) 
is also targeted by commercial fishers using a variety 
of gear, including traps, mesh and haul nets and 
electrofishing. 

There are a number of options to prevent or 
minimise mortality of air-breathing wildlife species 
in traps. The most direct way is to ban fishing in areas 
where these potentially vulnerable species occur. 



REDUCING BY-CATCH OF PLATYPUS 





UNREGULATED 






FISHERY 




,r 




w 


PREVENT 




MINIMISE 


BY-CATCH 




BY-CATCH 






, r 


REGULATE 


REGULATE 


TO CLOSE 




FISHING 


WATERS 




METHODS 



TRAP 

MODIFICATION 

NECESSARY 



BY-CATCH 
REDUCTION 

DEVICE 

MINIMISES 

BY-CATCH 

MORTALITY 



BY-CATCH 

REDUCTION 

DEVICE NOT 

APPROPRIATE 

ORIS 
INEFFECTIVE 



STANDARD 

TRAP 

MINIMISES 

BY-CATCH 

MORTALITY 



MODIFY TRAP 
TO PERMIT 

NON-TARGET 

SPECIES TO 

ESCAPE 



MODIFY TRAP 

TO PROVIDE 

NON-TARGET 

SPECIES WITH 

AIRSPACE 



Figure 1. Schematic diagram of possible options available to achieve by-catch reduction of air-breathing 
species in fisheries operations. 



However, maintaining a commercial fishery, while still 
addressing the issue of by-catch mortality, is to adopt 
capture methods which minimise by-catch. Mortality 
of air-breathing non-target species can be reduced or 
prevented by trap modifications, such as fitting devices 



to keep non-target species out (By-catch Reduction 
Device - BRD), providing a route to let them escape 
or permitting access to an airspace once they have 
entered a trap. Figure 1 summarises these possible 
options, which need to be explored in relation to the 
following issues: 



260 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 



Fishery requirements . The practicalities and economics 
of the fishery, in terms of trap design and cost, catch 
per unit effort, size of target species, and even the 
necessity to hide traps from possible interference and/ 
or vandalism must be considered. For instance, a device 
which reduces by-catch but unduly restricts the entry 
of the target species into a trap may be economically 
unviable. 

Behaviour of target species . It is necessary to know 
the reactions of the target species to trap modifications 
provided for non-target species. For example, the target 
species may escape via holes provided for the non- 
target species, or its behaviour could prevent the non- 
target species from utilising air spaces or escape routes 
provided. 

Behaviour of non-target species . In fishing areas where 
a number of potential by-catch species occur, escape 
holes, BRDs or air spaces in traps may not be suitable 
for all potential non-target species. For example one 
species may use an escape hole in a trap which will 
not be used by another species. 

This paper reviews past efforts to reduce the 
mortality of platypuses in the eel, yabby and carp 
fisheries and reports on a number of recent studies 
carried out to assess the effectiveness of trap 
modifications designed to reduce by-catch mortality 
of this species in these fisheries. The three fisheries 
are reviewed in separate sections of the paper and the 
experiments pertinent to each are discussed within 
these sections. 



THE EEL FISHERY IN NEW SOUTH WALES 

Freshwater eels were initially captured in 
upper estuarine waters of NSW as a by-catch of other 
fisheries. A fledgling industry targeting eels, based on 
the use of traps, was established in the early 1980s. At 
that time prices for eels were low but in the late 1980s 
and early 1990s a high-value export market to Asia 
was established. This increased interest in the fishery 
and the adoption of potentially more productive fishing 
methods. Requests were made by fishers to extend their 
operations into freshwaters using fyke nets (Figure 2a), 
which were known to be involved in the mortality of 
air-breathing wildlife species in the eel fisheries both 
in Tasmania and in Victoria (Jackson 1979; Beumer 
et al. 1981; Grant 1991). The potential fishers drew 
attention to a brief experiment in Lake Crescent and 
Dee Lagoon in Tasmania, where two fyke nets 
screened with 100 mm square mesh grids, and two 
unscreened control nets, were deployed in those lakes 
for six days. During that time, two platypuses were 
captured in the unscreened nets but none were captured 



in the ones with the screens in place (Grant 1991). 
While it appeared from this very limited experiment 
that a 100 mm mesh screen may have been effective 
in reducing platypus by-catch in Tasmania, an 
experiment done in the upper Shoalhaven River did 
not support this contention (Grant, unpublished data). 
Six platypuses (two female and four male) were placed 
separately between the river bank and the wing of a 
fyke net with a 100 mm mesh entrance screen in place. 
Two of these animals moved off after bumping the 
mesh and did not enter the fyke net but the other four 
either passed straight through into the net, or did so 
after first investigating the screen. 

At the time it was also known that elevating 
the cod end of fyke nets above the surface was effective 
in permitting platypuses to breathe and survive capture 
(Jackson 1979; Beumer etal. 1981;Grant 1991;Figure 
2b). Unfortunately professional fishers were 
unprepared to do this, as they feared their catch could 
be stolen and/or their equipment vandalised if it was 
visible above the surface. 

As a result of the brief experiment with the 
Shoalhaven River platypuses described above, and 
advice from experts in the other states regarding the 
poor compliance of fishers to fit BRDs and/or to raise 
the cod-ends of their nets above the water level, the 
request by fishers to use fyke nets for eels, and to extend 
the fishery to freshwater streams was denied by NSW 
Fisheries. Instead, the fishery was restricted to estuarine 
waters, a limited number of impoundments and private 
farm dams, using baited traps without wings to direct 
animals Into the traps (NSW Fisheries Eel Policy 
Document, May 1992). 

The standard eel traps used in the fishery are 
shown in Figure 2c. They consist of a metal rod frame 
50 cm wide by 40 cm high by 90 cm long covered 
with 30 mm mesh polyethylene netting. The single 
entrance funnel (or 'valve') is located in one end of 
the trap. The opening in the funnel consists of a hole 
in the netting stretched firmly into a 100 mm wide 
slot, and pulled approximately 20 cm into the trap. 
The traps used in estuaries have a 1.5 m long cod end 
(bag with a draw-string) on the opposite end of the 
trap from the entrance funnel. Those used in freshwater 
impoundments and farm dams are similar to the estuary 
trap, but have a 5 m long cod end. A 150-200 mm 
diameter float is fastened inside the cod end near the 
draw-string and from one to three 50 cm diameter 
aluminium hoops are fastened to the inside of the cod 
end to keep the passage to the surface open. These 
traps are normally baited with frozen pilchards or 
mullet to attract eels. 

In the late 1990s anecdotal reports to the 
National Parks and Wildlife Service, NSW Fisheries 
and one of the authors (TRG) indicated that platypuses 



Proc. Linn. Soc. N.S.W., 125, 2004 



261 



REDUCING BY-CATCH OF PLATYPUS 



a 



cod end 




hoops 



^■^^. ^ ■^^iiC ^ i^^ V ^Jc V -^Ai^JWi'ci^Jc ) L-JCJC ^ -^iiJc\» l C\. ^ ^.Jc l ».^\. ^ JCJc l C 




entrance funnel 



Figure 2. (a) Fyke net used in eel Fisheries in Tasmania and Victoria, (b) Commercial eel trap used in the 
impoundment or farm dam eel Fishery, showing the elevated cod-end creating an air space, (c) Typical eel 
trap used in the tidal estuary Fishery in New South Wales. Note the entrance funnel or 'valve' which 
permits animals to enter the traps in one direction (wide outside to narrow inside). Animals are unable to 
locate the narrow inside entrance to escape. In Experiment 1 grids were placed at the narrow end of the 
funnel and in Experiment 2 at the wide end. 



262 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 



were being drowned in eel traps, not only in the upper 
reaches of some estuaries (where tidal influence 
changed with river discharges) but also in farm dams 
and impoundments (where air spaces were not 
consistently being maintained in the cod ends of traps). 
As a result, the following experiments were undertaken 
to determine if it was possible to reduce this mortality 
of platypuses by trap modification. 

EXPERIMENT 1 - Investigation of grid sizes for a 
platypus exclusion device 

The objective of this experiment was to 
determine the optimum grid size for excluding most 
platypuses from eel traps. The experiment was 
conducted in two pools on the Wingecarribee River in 
New South Wales from 17-19 February 2000. 

Methods 

The entrance funnels in eight standard eel 
traps were fitted with grids of different sizes. Each 
grid was a square divided into four equal openings; 
the openings in these grids ranged from 55 to 90 mm, 
in 5 mm increments. The plastic material used to make 
the grids was reinforced with lengths of 3 mm wire. 
The traps were fastened end to end (in order of 
decreasing grid size) and placed on a flat sandy area 
in the pools where platypuses were to be captured 
(Figure 3a). Water depth varied between traps but all 
had an airspace to allow the platypuses to breathe 
during the experiment. 

Trials were done in different pools on two 
days. Platypuses were captured using unweighted gill 
nets (Grant and Carrick 1974) during the evening or 
morning. Once the required numbers of platypuses 
were captured, each individual was measured and 
weighed, then tested individually in the experiment. 
Platypuses were placed through an access door into 
the first trap leading into an entrance funnel with the 
90 mm grid in place (Fig. 3a). Red-filtered lights were 
used to observe the animals at night, as observations 
in captivity indicated that platypuses are less 
responsive to disturbance under red light illumination 
(Grant, personal observation). The time that animals 
remained in each trap before passing through each grid 
was recorded, along with the number of attempts that 
each animal made to pass through the entrance funnel 
into the next trap in the series. Animals were removed 
from the experiment and released immediately if they 
remained in any trap for more than 15 minutes. 

Results 

A total of ten platypuses were used in the 
trials, comprising two adult males (1 190 and 1760 g), 



six adult females (890-1060 g) and two juvenile 
females (700 and 760 g). Data are summarised in Table 
1. 

Trial 1 : Animals tested at night were reluctant 
to pass through the 85 mm grid and none passed 
through the 75 mm grid, while a single female captured 
in the morning, and tested in daylight readily, passed 
through all grid sizes, although exhibiting some delay 
at the 80 and 70 mm grids. However, it was noted that 
the traps with 85-70 mm grids, which were apparently 
difficult for the animals to negotiate, were located in 
slightly shallower water than the rest of the traps. The 
water level in these traps was located at or just above 
the top of the grid, whereas the water level in the other 
traps was well above the top of the entrance grids. It 
was thought that this difference in water depth may 
have influenced platypus behaviour. Subsequently, all 
traps were placed in deeper water (well over the top of 
the grid) during the second trial. 

Trial 2: The largest male (1760 g) could not 
pass through the 65 mm grid, but the smallest female 
(700 g) passed through each grid in less than 1 minute. 
The two slightly larger females did not initially pass 
through the 55 mm grid. However, it was found that, 
due to some unevenness on the bottom of the pool, the 
trap with this grid was in slightly shallower water than 
the preceding traps in the series. After moving this 
last trap to a position in slightly deeper water, animals 
passed through the 55 mm grid almost immediately. 

The data from Trial 1 indicated that there was 
a greater reluctance for platypuses to negotiate the grids 
when the traps were less submerged. However, Trial 
2 confirmed that female platypuses of up to 1 kilogram 
in weight could pass through a 55 mm grid. Animals 
smaller than 1 kg passed through easily, while the 1 
kg female had a tighter squeeze. Only one male 
platypus was captured for use in Trial 2. This was the 
largest animal tested (1760 g) and was stopped by the 
65 mm grid. A grid between 55 and 65 mm would 
apparently be required to exclude most adult male 
platypuses. 

EXPERIMENT 2 - Investigation of possible 
avoidance of entrance grids by free-swimming 
platypuses 

In Experiment 1, each platypus was closely 
confined inside the traps so there was an imperative to 
find an escape route. However, two of the four animals 
in Trial 2 hesitated, and made more than one attempt 
to pass through the 70 mm grid, indicating possible 
deterrent effect of this grid size. Experiment 2 was 
designed to test whether grids across the outer end of 
the entrance funnel (Figure 3b) deterred foraging 



Proc. Linn. Soc. N.S.W., 125, 2004 



263 



REDUCING BY-CATCH OF PLATYPUS 



entrance funnels 




water level 




netting 
enclosure 



Figure 3. Set up used in Experiments 1 and 2 to test the effectiveness of by-catch reduction devices (BRDs) 
on entry of platypuses into eel traps. (Top) Experiment 1. Traps were attached together in a line with 
grids of different sizes at the narrow end of each entrance funnel or 'valve'. (Bottom) Experiment 2. 
Mesh enclosure in a river pool with trap entrance attached. Note the position of the replaceable rectangular 
grid across the outer (wide) entrance of the funnel. 



platypuses from entering traps. The experiment was 
done in a pool on the upper Shoalhaven River in the 
southern tablelands of New South Wales from 17-19 
March 2000. 

Methods 

A circular enclosure, 1.5 m high x 3 m 
diameter, made from 1 mm mesh monofilament gill 
net material, was constructed in a pool between the 
two netting sites where platypuses were captured for 
the experiment. The enclosure was designed so that 
the only possible escape for a platypus was through 
the grid of the entrance funnel of a trap inserted in the 
enclosure wall. Square grids, made from 4 mm steel 
rods, with 50, 60, 70 and 80 mm openings were used 
in this experiment. Each trial was done by attaching a 



replaceable grid to the entrance funnel of the trap, then 
placing a platypus into the enclosure (Figure 3b). At 
night, red-filtered lights were used to observe the 
animals. The time each animal remained in the 
enclosure before passing through the grid was 
recorded, along with the number of attempts that each 
made to pass through the grid. If an animal did not 
pass through a particular grid in the test series, this 
was replaced by the next larger grid in the series and 
the observations repeated. After the first animal was 
obviously unable to exit the 50 mm grid, the trials on 
all others were begun with either the 60 or 70 mm 
grid. 



Results 



Eight relatively small platypuses (ranging in 



264 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 



Table 1. Details of platypuses exiting through the various grid sizes within the funnels of 
eel traps in the two trials of Experiment 1. + = animal exited specific grid size; 
X = platypus did not exit through specific grid size. 



Sex/ Weight 90 mm 85 mm 80 mm 75 mm 70 mm 65 mm 60 mm 55 mm 

age (g) Grid Grid Grid Grid Grid Grid Grid Grid 



X 



Trial 1 






Male 






Adult 


1190 


+ 


Female 






Adult 


1060 


+ 


Female 






Adult 


1030 


+ 


Female 
Adult 


1020 


+ 


Female 
Adult 


920 


+ 


Female 
Adult 


890 


+ 



X 



X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


+ 


X 


X 


X 


X 


X 


+ 


+ 


+ 


+ 


+ 


+ 


+ 


X 


X 


X 


X 


X 



Exited 



6/6 



4/6 



3/6 



1/6 



1/6 



1/6 



1/6 



1/6 



Trial 2 




Male Adult 


1760 


Female 




Adult 


1000 


Female 




Juvenile 


760 


Female 




Juvenile 


700 



X 



X 



X 

+ 
X 

+ 



Exited 



4/4 



4/4 



4/4 



4/4 



4/4 



3/4 



3/4 



2/4 



size from 500 to 940 g) were tested in the enclosure at 
night. Results of the grid-deterrent trials are shown in 
Table 2. 

The first platypus was initially placed in the 
enclosure with the 50 mm grid. After six attempts to 
go through the grid it was apparent that the animal 
would not fit through the spaces. After several tentative 
attempts at the 60 mm grid it appeared to stop trying 
to escape through the subsequent grids and remained 
in the enclosure even after the largest grid was 
completely removed. The test with the second platypus 
was started with the 60 mm grid in place, but this 
platypus was less active than the first animal and made 
only one tentative attempt to pass through this grid. It 
then readily passed through the 70 mm grid after only 
one attempt. Trials with the next three platypuses were 
all started with the 60 mm grid. All three of these 
animals swam past the grid at least once before 
escaping through it. The last three animals were 
initially trialed with the 70 mm grid, and all passed 



through it at the first attempt. Overall, two animals 
out of five appeared to be deterred by a 60 mm grid 
(40%) and only a single animal was deterred by a 70 
mm grid (Table 2). 

EXPERIMENT 3 - Platypus behaviour in the 
elevated cod ends of traps modified for use in farm 
dams and impoundments 

The objective of this experiment was to record 
the behaviour of platypuses in modified eel traps used 
in impoundments and farm dams (Figure 2c) and to 
investigate their ability to negotiate the long cod end 
extension to the air space. The experiment was done 
in a pool on the upper Shoalhaven River from 17-19 
March 2000. 

Method 

Two impoundment eel traps, with 5 m cod 
ends (Figure 2c) were placed in a pool of 0.5 m depth. 



Proc. Linn. Soc. N.S.W., 125, 2004 



265 



REDUCING BY-CATCH OF PLATYPUS 



Table 2. Details of platypuses deterred from entering the 
various grid sizes across the entrances of eel traps in 
Experiment 2. Animals are arranged in the order in which 
they were used in the experiment. + = animal passed through 
specific grid size; X = platypus did not pass through specific 
grid size i.e. deterred; - no data; 



Sex/Age 


Weig 


ht 50 mm 


i 60 nun 


l 70 mm 80 mm 




(g) 


Grid 


Grid 


Grid Grid 


Female 










Adult • 


800 


X 


X 


X X 


Female 










Juvenile 


500 


- 


X 


+ 


Male 










Juvenile 


800 


- 


+ 


- 


Male 










Juvenile 


740 


- 


+ 


- 


Male 










Juvenile 


640 


- 


+ 


- 


Female 










Adult 


940 


- 


- 


+ 


Female 










Adult 


900 


- 


- 


+ 


Female 










Juvenile 


690 


- 


- 


+ 



One trap had three evenly spaced hoops in the cod end and the 
other had only one hoop near the airspace. The cod end of each 
trap was stretched and tied off above the surface of the water to 
a star-picket. Three platypuses (one male and two females) were 
placed consecutively in the trap with three hoops, and one 
female platypus was placed in the trap with one hoop. Each 
platypus was observed for 15 to 20 minutes before being 
released. 

Results 

In each case the platypus spent several minutes searching the 
inside of the trap before travelling up the cod end to the airspace. 
Each took several breaths then travelled to the trap where it 
again searched around or 'wedged' itself under the entrance 
funnel. Within five to eight minutes each would again travel 
up to the airspace for several breaths before returning to the 
trap. Platypuses travelled back and forth from the trap to the 
airspace 2-3 times during the 15-20 minutes they were confined 
in the trap. 

DISCUSSION - Eel Trap Experiments 

The results of Experiments 1 and 2 indicated that a 
grid of 50-55 mm would be necessary to exclude platypuses 
from entry into eel traps. Such a by-catch reduction device 
(BRD) would almost certainly affect the catch rates and sizes 
of eels (Koed and Dieperink 1 999). This would be unacceptable 



to commercial fishers, particularly those 
fishing for adults of the long-finned species 
(Anguilla reinhardtii). Free-ranging 
platypuses may be deterred from entering 
traps fitted with external grids of 70 mm or 
less across the entrance funnels but such 
screening would be unlikely to significantly 
reduce platypus by-catch in eel traps. 

Raising the cod end to provide an 
air space would facilitate the survival of 
platypuses captured in eel traps fitted with 
elongated cod ends. Platypuses captured in 
these traps were reluctant to stay at the 
surface and preferred to remain submerged 
in the trap between taking breaths. This 
behaviour, which minimises the time spent 
at the surface, may be a mechanism to avoid 
natural predation. Because platypuses must 
breathe at least every 2-10 minutes (Bethge 
2002), captured individuals would need to 
travel back and forth to the airspace many 
times during any extended period of 
confinement after capture. This would be 
stressful and energetically demanding. It is 
essential that captured animals be released 
as soon as possible after capture. Studies 
using fyke nets (with elevated cod ends) to 
capture fish have shown that platypuses can 
survive for periods of up to 24 hours (Grant 
and NSW Fisheries, unpublished data). 
However, hypothermia has been reported in 
platypuses restrained in fyke nets after a few 
hours in cold conditions (Serena, personal 
communication). The current regulations in 
New South Wales demand that eel traps be 
inspected at least every 24 hours. 

The platypus forages aerobically 
for short periods by holding its breath, 
following a comparatively large inspiration 
of air after each dive (Evans et al. 1994; 
Bethge 2002). The behaviour observed in 
this study of 'wedging' themselves under an 
object, and reducing energetic demands by 
remaining stationary, has been reported in 
captivity to last up to 11 minutes (Evans et 
al. 1994; Bethge et al. 2001; Bethge 2002). 
The function of this behaviour and its 
occurrence in the wild has not been 
determined. However, from the perspective 
of by-catch mortality this behaviour would 
not prevent platypuses from being drowned 
in completely submerged traps during 
normal fishing operations, which demand a 
period of trap submergence of hours rather 



266 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 



than minutes. 

Observation of traps with airspaces 
maintained only by the use of a float has shown that 
the cod end can easily become twisted or bunched. 
This situation would undoubtedly prevent a captured 
air-breathing species from reaching the airspace. This 
can be avoided by stretching the cod end tightly to a 
fixed point, either on the bank or a star picket driven 
into the bottom of the water body. It should be noted 
however, where traps are set with elevated cod ends 
attached to a fixed point, allowance needs to be made 
for anticipated rises in water level as a result of rainfall 
and/or tidal influences. Attachment of the cod ends of 
eel traps to a fixed point is mandatory under regulations 
for the use of eel traps in impoundments and farm dams 
in NSW. 



THE COMMERCIAL AND RECREATIONAL 
YABBY FISHERY IN NEW SOUTH WALES 

The results of the experiments done to 
evaluate the effectiveness of devices to prevent or deter 
platypuses from entering eel traps are also directly 
applicable to both the commercial and recreational 
'yabby' [freshwater crayfish] fisheries. Based on the 
lack of adverse reports and on the assumption that the 
traps used to capture yabbies were small and did not 
have mesh wings to direct foraging platypuses into 
them, Grant (1993) suggested that "yabby fishing poses 
little threat to platypuses". This conclusion is now 
thought to be incorrect, as anecdotal reports from a 
number of states suggest that yabby traps were 
affecting some local platypus populations. These traps 
have also been implicated in the mortality of other non- 
target species, especially freshwater turtles. The 
drowning of as many as five platypuses in a single 
yabby trap has been reported, although the species' 
attraction to these traps is not fully understood. 
Platypuses are known to locate their prey by sensing 
the electrical fields generated by muscular activity of 
the prey species, especially large food items such as 
yabbies (Pettigrew et al. 1998). A trap containing live 
yabbies may therefore attract platypuses during their 
normal foraging activities. Once there is a dead 
platypus in a trap, more yabbies may feed on the 
decomposing carcass, which could in turn attract other 
platypuses into the trap. 

Rankin (2000) suggested that a fixed ring 60- 
70 mm in diameter may prevent platypuses from 
entering traps and also facilitate their escape. Some 
commercially available yabby traps are fitted with 90 
mm entrance rings, which are effective in excluding 
larger turtles but which are still reported to have 



drowned platypuses. The experiments described above 
for eel traps indicate that a 90 mm diameter ring is too 
large to exclude platypuses. Similarly, neither the 
experiments reported here nor anecdotal observations 
support Rankin's (2000) suggestion that platypuses 
could escape by returning through a fixed entrance 
ring. 

Allanson and Thurstan (1999) evaluated the 
effect of entrance rings of different diameters in yabby 
traps using relatively small captive-bred yabbies 
(Cherax destructor). These trials showed that the 
smallest ring tested (63 mm) still permitted yabbies of 
the same size to enter the experimental traps as were 
entering the control traps with no rings fitted. However, 
the experimental traps caught substantially fewer 
yabbies. When the results of Allanson and Thurston's 
(1999) experiments were discussed with commercial 
fishers, it was concluded that the use of such a small 
entrance ring was not a viable option for the 
commercial yabby fishery. 

Current regulations in New South Wales 
exclude the use of traps in commercial and recreational 
yabby fishing from known platypus waters and 90 mm 
rings are required in all yabby traps to exclude most 
turtles. Closed waters are located east of the Newell 
Highway, from the Victorian border (Murray River) 
to the Queensland border (Macintyre River), along with 
local closures around Deniliquin on the Edward River, 
Echuca on the Murray River and between Narrandera 
and Darlington Point on the Murrumbidgee River, 
where platypuses are also know to occur. 



THE CARP FISHERY IN NEW SOUTH WALES 

Carp (Cyprinus carpio) were probably first 
introduced into Australia around 1850 but did not 
spread until the introduction of the 'Boolarra' strain 
in the 1960s. Ecological effects of high densities of 
carp are poorly understood, but increased bank 
damage, disturbance of aquatic macrophytes and 
turbidity are all possible consequences. The overall 
disruption of riverine food webs by the large biomass 
of carp is thought to be detrimental to freshwater 
ecosystems (Schiller and Harris 2001). Carp are 
harvested in New South Wales using a variety of gear, 
including traps, haul and mesh nets, and electrofishing 
equipment. There is considerable overlap between the 
distribution of carp and platypuses (Boulton and Brock 
1999), making the use of submerged traps a concern 
in this fishery. 

A drum trap was constructed by NSW 
Fisheries (Fig. 4), which was designed to permit the 
escape of air-breathing vertebrate species, including 



Proc. Linn. Soc. N.S.W., 125, 2004 



267 



REDUCING BY-CATCH OF PLATYPUS 



platform 



platypus escape 
holes 





turtle escape hole 



entrance funnel 



escape hole 




entrance funnel 

Figure 4. (Top) Modified drum trap showing escape hole in the roof 
above the mesh platform. Note the entrance funnel (or 'valve') on the 
left end of the drum. The entrance was sealed in the experiments and 
the triangular escape holes were made at the base of this funnel. 
(Bottom) Inside the trap showing the position of the steel mesh platform 
below the escape opening. 



platypuses, water rats, turtles and diving birds, through 
a hole in the trap's roof. A wire-mesh platform was 
positioned below the escape hole so that small 



vertebrate species could pass 
through the 8 cm gap between it 
and the roof of the trap and exit 
through the escape hole, while 
larger carp would not be able to 
escape. Carp are also inclined to 
congregate near the bottom of a 
trap. The design assumed that air- 
breathing species would tend to 
swim towards the surface and 
search along the roof of the trap 
for a means of escape (surface/ 
search behaviour). The objective 
of the following experiment was 
to test the effectiveness of the 
escape device for platypuses. 

EXPERIMENT 4 - Assessment 
of escape of platypuses from a 
prototype carp trap 

Platypuses close their 
eyes, ears and nostrils when under 
water, using the sensory 
mechanisms in their bills to find 
their way around (Pettigrew et al. 
1998). It was expected that 
platypuses in the experiment 
would exhibit surface/search 
behaviour and be able to escape 
from the modified drum trap. The 
experiment was done in several 
pools on the Wingecarribee 
River, New South Wales from 25- 
27 November 2002 to determine 
if this expectation was realised. 



Method 

The trap consisted of a 
90 cm diameter x 170 cm long 
cylinder, covered with black 
plastic mesh (55 mm x 40 mm), 
except at the entrance end, where 
a conical funnel or 'valve' made 
from 3 mm diameter braided 
polyethylene trawl netting was 
strung tightly between the 
circular steel frame at one end of 
the trap and an oval ring rigidly 
suspended inside the trap (Figure 
4). 

The trap was fully 
submerged in the pools from which the platypuses were 
captured. The trap was oriented with the escape hole 
uppermost. A remote lens for a video camera was 



268 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 



Table 3. Results of Experiment 4. Assessment of escape of platypuses from the carp trap in the 
Wingecarribee River. 



Sex/Age Weight (g) Length Time in 

(cm) trap (sec) 



Approaches Escape 
to platform 



Adult 


1080 


Male 




Adult 


1880 


Male 




Juvenile 


1790 


Male 




Adult 


1880 


Male 




Juvenile 


1400 



48.5 
55.2 
56.5 
57.5 
53.0 



150 
165 
180 
180 
30 



No 
No 
No 
No 
Yes 



mounted inside the trap to record the behaviour of the 
animals and these images were stored for later analysis. 
Platypuses were captured using unweighted 
gill nets (Grant and Carrick 1974). Each animal was 
weighed and measured, then temporarily marked with 
a piece of brightly coloured tape attached to the tail, 
making the platypuses more visible to observers and 
to the video camera. Based on observations reported 
above and Bethge (2002), who reported a maximum 
foraging dive duration of 138 seconds, individuals 
were immersed for a maximum of 3 minutes before 
the trap was lifted to permit them to breathe. If they 
exited the trap prior to lifting, the elapsed time was 
recorded. The numbers of times each animal 
approached the platform below the escape hole was 
recorded. All animals were used only once in the 
experiment and remained in the trap for no more than 
3 minutes. 

Results 

Table 3 shows the dimensions of the 
platypuses used, the time in the trap, the number of 
approaches to the platform below the escape hole, and 
whether or not individuals escaped. Only one juvenile 
male platypus managed to find the escape hole (after 
30 seconds in the trap), but showed reluctance to leave 
the steel ring around the hole. It re-entered the body 
of the trap three more times before finally leaving the 
trap completely. This animal repeatedly relocated the 
escape hole after re-entering the trap, taking 30, 50 
and 50 seconds respectively, before finally escaping. 
The other four trial animals failed to find the escape 
hole and were released after 2-3 minutes. 

Contrary to expectation, platypuses 
(including the one which escaped) spent most of the 
time investigating the bottom or ends of the trap, rather 
than exhibiting surface/search behaviour. In fact, they 
seemed to actively avoid the platform area below the 



escape hole. All animals searched with their bills 
around the corners of the trap between the sides and 
ends. The video showed them frequently investigating 
the acute angled edge between the base of the entrance 
funnel and the sides of the trap. When released, all 
animals were observed to surface and appeared to be 
breathing heavily. 

EXPERIMENT 5. Assessment of escape of 
platypuses from the modified carp traps 

In Experiment 4, platypuses were observed 
continually searching for an escape hole around the 
corners of the trap. It was therefore decided to test the 
effectiveness of escape holes positioned around the 
base of the entrance funnel. Because the gap between 
the funnel and the sides of the trap was quite narrow at 
the base of the funnel, it was considered that most carp 
would be too large to access openings in this position. 
Experiment 5 tested the effectiveness of these 
modifications. The experiment was done in one pool 
on the Wingecarribee River on 27 November 2002 and 
then in four pools on the upper Shoalhaven River from 
21-23 December 2002. 

Methods 

Every third mesh attached to the trap frame 
at the base of the funnel was released and tied back to 
provide 90 x 90 x 90 mm triangular openings (Fig. 4, 
top). In the initial trial in the Wingecarribee River these 
openings were made only in the upper half of the trap, 
but in the later trials in the upper Shoalhaven River, 
openings were made in both the upper and lower halves 
of the trap. 

Fourteen platypuses were individually placed 
in the submerged trap as described in Experiment 4. 
Again observations were made of the number of times 
animals approached the platform below the escape 



Proc. Linn. Soc. N.S.W., 125, 2004 



269 



REDUCING BY-CATCH OF PLATYPUS 



Table 4. Results of Experiment 5. Assessment of escape of platypuses from the modified carp trap. * not 
observed escaping but were not present in trap when it was lifted after 3 minutes; - escape holes only 
available in upper part of this trap. 



Sex 


Weight 


Length 


Time in 


Approaches 


Escape 


Escape 




(g) 


(cm) 


trap (sec) 


to 


platform 




location 


Female 


850 


43.0 


85 


1 




Yes 




Female 


690 


41.0 


15 







Yes 


lower 


Female 


900 


46.0 


22 







Yes 


lower 


Female 


940 


43.5 


15 







Yes 


lower 


Female 


900 


44.0 


40 







Yes 


upper 


Female 


790 


41.0 


41 







Yes 


upper 


Female 


870 


43.5 


140 







Yes 


upper 


Female 


930 


44.0 


33 







Yes 


upper 


Female 


860 


44.0 


<180* 







Yes 


lower 


Female 


840 


43.5 


45 







Yes 


upper 


Female 


790 


43.0 


156 







Yes 


upper 


Male 


1850 


55.2 


35 


1 




Yes 


- 


Male 


1740 


52.0 


<180* 







Yes 


lower 



hole. Escapes through the triangular holes at the base 
of the entrance funnel were partitioned as being from 
the 'upper' or 'lower' openings in the trap. Some 
underwater video observations were made but the 
turbidity of the pools made viewing difficult. However, 
brightly coloured tape attached to the tails of the 
animals (see Experiment 4) usually permitted their 
movements in the trap to be observed. Again, if the 
platypus was not seen to escape, the trap was lifted 
from the water after a maximum of 3 minutes. 

Results 

Thirteen platypuses escaped from the 
openings around the base of the entrance funnel of the 
trap within 3 minutes (Table 4). As was observed in 
Experiment 4, all animals attempted to find an escape 
route around the bottom or ends of the trap. Another 
individual used in the initial trial located a hole 
inadvertently left at the bottom of the trap (which was 
sealed before subsequent trials). No preference was 
shown for escape location, with six animals exiting 
from the 'upper' and 5 from the 'lower' openings, 
where both were available (Table 4). One individual 
moved into the space between the platform and the 
escape hole but did not find the hole, submerging again 
and leaving the trap by one of the openings at the base 
of the entrance funnel. Only two individuals 
approached the platform at any time during their 
confinement in the trap. In two instances the platypuses 
could not be seen, but were no longer in the traps when 
they were lifted after 3 minutes. It was presumed that 
they had exited the lower holes, as they were not seen 
leaving the upper ones, which were visible to the 
observers. 



DISCUSSION - Carp trap experiments 

Experiment 4 indicated that the unmodified 
carp trap would probably result in significant mortality 
of platypuses if deployed in areas where their 
distribution overlaps that of carp. However, experiment 
5 indicated that carp traps with appropriate escape holes 
could be used to reduce by-catch of platypuses. 
Platypuses over a size range of 690-1880 grams were 
able to exit quite quickly (15-156 seconds) through 
the 90 mm triangular openings in the modified carp 
trap. 

It should be noted that the platypuses used in 
these experiments were not particularly large. There 
is considerable sexual dimorphism in the species, with 
the average male being around 75% heavier and 20% 
longer than females (Carrick 1995; Grant 1995; 
Connolly and Obendorf 1998). Individuals of up to 
twice the size of those used in current experiments are 
found in some mainland areas (especially west of the 
Great Dividing Ranges; Carrick 1995; Grant 1995) and 
in Tasmania males may reach up to three kg (Connolly 
and Obendorf 1998). Further experiments are required 
to determine the size of escape holes effective for larger 
platypuses. In the interim, the authors recommend 
triangular openings of 100 x 100 mm for east-flowing 
streams in New South Wales and openings of at least 
120 x 120 mm for west-flowing streams in the state. 
Trials would also need to be carried out to assess the 
effectiveness of retaining captured carp in the presence 
larger escape holes. 

The unexpected lack of surface/search 
behaviour in platypuses during Experiments 4 and 5 



270 



Proc. Linn. Soc. N.S.W., 125, 2004 



T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 



indicates the importance of field trials of fishing 
equipment with regard to specific wildlife species. The 
reason for the unexpected lack of surface/search 
behaviour in water can only be speculated upon. 
Platypuses frequently forage among dense woody 
debris and under submerged overhanging banks (Grant 
1995 and personal observation). It may be that a 
behavioural response of moving down and/or sideways 
away from an obstruction during foraging may be of 
greater survival value than attempting to rise directly 
to the surface when seeking an escape route. No 
'wedging' behaviour (Evans et al. 1994; Bethge et al. 
2001; Bethge 2002; Experiment 3) was exhibited by 
animals in the carp traps. Rather, all individuals 
searched constantly for an escape route. 



GENERAL CONCLUSIONS 

The results of the literature reviewed and 
experiments presented in this paper indicate that any 
fishery in freshwaters of New South Wales based on 
the use of traps should not be operated as an 
unregulated fishery (Figure 1) if reducing platypus 
mortality is a priority. By-catch minimisation has been 
possible in the eel fishery by a combination of closures 
of some inland waters and by modifications to provide 
an airspace in traps used in farm dams and 
impoundments. Exclusion devices (e.g. grids across 
the entrance funnels of traps) do not provide a 
commercially viable option for reducing the by-catch 
of platypuses in eel or yabby traps. Banning of yabby 
traps from areas where platypuses occur is currently 
the only available means of avoiding by-catch 
mortality in this fishery. The commercial and 
recreational yabby fisheries in New South Wales are 
currently restricted to waters where platypuses do not 
commonly occur or are very uncommonly reported. 
Trap modifications, which permit the escape of 
platypuses, appear to be the most feasible means of 
by-catch minimisation in the use of traps to capture 
carp. 



ACKNOWLEDGMENTS 

This work was conducted under Animal Research 
Authorities (ACEC 99/13 and ACEC 02/12) from the NSW 
Fisheries Animal Care and Ethics Committee and under NSW 
Fisheries Scientific Research Permit F84/1245 (TRG) and 
NSW National Parks and Wildlife Service Scientific 
Investigation Licence A 1 84 (TRG). Frank Jordan constructed 
the grids used in Experiment 2 and Stuart Scott and the Izzard 
and Laurie families allowed us to work on their properties 
on the Wingecarribee and Shoalhaven Rivers. Dr Melody 



Serena, of the Australian Platypus Conservancy is 
acknowledged for her personal communication regarding 
hypothermia in platypuses captured in fyke nets. Funding 
for much of the reported work was provided by NSW 
Fisheries. The authors thank Mike Augee and an anonymous 
referee for their valuable comments and suggestions. 



REFERENCES 

Allanson, M. and Thurstan, S. (1999). 'Yabby {Cherax 
destructor) trap by-catch reduction.' 
Unpublished report NSW Fisheries-Inland 
Management. August/September, 1999. 

Bethge, P. (2002). Energetics and foraging behaviour of 
the platypus Ornithorhynchus anatinus. PhD 
thesis. University of Tasmania, Hobart. 

Bethge, P., Munks, S. and Nicol, S. (2001). Energetics 
and locomotion in the platypus, 
Ornithorhynchus anatinus. Journal of 
Comparative Physiology - B, Biochemical, 
Systematic and Environmental Physiology 171, 
497-506. 

Beumer, J.P., Burbury, M.E. and Harrington, D.J. (1981). 
The capture of fauna other than fishes in eel and 
mesh nets. Australian Wildlife Research 8, 673- 
677 

Boulton, A.J. and Brock, M.A. (1999). 'Australian 
Freshwater Ecology. Processes and 
Management'. Gleneagles Publishing, Glen 
Osmond, South Australia. 

Carrick, F.N. (1995). Family Ornithorhynchidae. 

Platypus. In 'The mammals of Australia' (Ed R. 
Strahan) pp. 35-38. (Reed Books: Chatswood). 

Connolly, J.H. and Obendorf, D.L. (1998). Distribution, 
captures and physical dimensions of the 
platypus {Ornithorhynchus anatinus) in 
Tasmania. Australian Mammalogy 20, 231-237. 

Evans, B.K., Jones, D.R., Baldwin, J. and Gabbott, G.RJ. 
(1994). Diving ability of the platypus. 
Australian Journal of Zoology 42, 17-27. 

Grant, T.R. (1991). The biology and management of the 
platypus {Ornithorhynchus anatinus) in New 
South Wales. Species Management Report # 5. 
NSW National Parks and Wildlife Service, 
Hurstville, New South Wales 

Grant, T.R. (1993). The past and present freshwater 

fishery in New South Wales and the distribution 
and status of the platypus, Ornithorhynchus 
anatinus. Australian Zoologist 29, 105-113. 

Grant, T. (1995). 'The platypus. Unique mammal'. 

University of New South Wales Press. Sydney. 

Grant, T.R. and Carrick, F.N. (1974). Capture and 
marking of the platypus, Ornithorhynchus 
anatinus, in the wild. Australian Zoologist. 18: 
133-135. 

Grant, T.R. and Denny, M.J.S. (1991). Distribution of the 
platypus in Australia with guidelines for 
management. Report to Australian National 



Proc. Linn. Soc. N.S.W., 125, 2004 



271 



REDUCING BY-CATCH OF PLATYPUS 



Parks and Wildlife Service. Mount King 
Ecological Surveys, Oberon, New South Wales. 

Jackson, P.D. (1979). Survey of fishes in the west branch 
of the Tarwin River above Berrys Creek. 
Victorian Naturalist 97, 11-14 

Koed, A. and Dieperink C. (1999). Otter guards in river 
fyke-net fisheries: effects on catches of eels and 
salmonids. Fisheries Management and Research. 
6: 63-69. 

Leadbitter, D. (2001). Bycatch Solutions. A Handbook for 
Fishers in Non-Trawl Fisheries. Oceanwatch 
and Fisheries Research and Development 
Corporation. Pyrmont, New South Wales, 
Australia. FRDC Project NO. 98201. 

Pettigrew, J.D., Manger, P.R., Fine, S.L.R. (1998). The 
sensory world of the platypus. Transactions of 
the Royal Society of London. Series B. 353, 
1199-1210. 

Rankin, T (2000). Status of the New South Wales 
commercial yabby fishery. NSW Fisheries 
Fishery Resource Assessment Series No. 9. 
Fisheries Research Institute, Cronulla. May 
2000. 

Schiller, C.B., Harris, J.H. (2001). Native and alien fish. 
In 'Rivers as Ecological Systems: the Murray- 
Darling Basin' (Ed W.J. Young) pp. 229-258. 
(Murray-Darling Basin Commission: Canberra). 

Serena, M. (2003). Wanted. Platypus sightings. Australian 
Society for Limnology Newsletter March 41, 36. 



272 Proc. Linn. Soc. N.S.W., 125, 2004 



Platypus Burrow Temperatures at a Subalpine Tasmanian Lake 

Philip Bethge 1 , Sarah Munks 2 , Helen Otley 2 , Stewart Nicol 1 

1 Division of Anatomy and Physiology, University of Tasmania, GPO Box 252-24, Hobart TAS 7001, 

Australia (Address for correspondence: Brandstwiete 19, 20457 Hamburg, Germany; philip@bethge.org '): 

2 School of Zoology, University of Tasmania, Private Bag 4,GPO, Hobart TAS 7001, Australia 



Bethge, P., Munks, S., Otley, H. and Nicol, S. (2004). Platypus burrow temperatures at a subalpine 
Tasmanian lake. Proceedings of the Linnean Society of New South Wales 125, 273-276. 

When platypuses are in their burrows, microhabitat is of great importance for energy conservation, especially 
where air temperatures frequently fall below freezing in winter. In this study, we investigated burrow 
temperatures of platypuses {Ornithorhynchus anatinus) living at a sub-alpine Tasmanian lake. Nine individual 
platypuses were equipped with time-depth recorders with integrated temperature sensors measuring ambient 
temperature. Burrow temperatures were recorded in two minute intervals for a total of 61 resting periods 
(duration: 5.45 to 27.20 hours) and were averaged over the period of resting. Mean burrow temperatures 
were 17.5 and 14.2°C (SD=2.76 and 0.89, respectively, n=9) in summer and winter, respectively, and 
ranged between 12.2 and 22.8°C for individual resting periods. In winter, burrow temperatures were held 
fairly constant over the resting period while in summer larger variations were observed. Burrow temperature 
in winter was found to be up to 18°C higher than outside air temperature. 

Manuscript received 26 November 2003, accepted for publication 8 January 2004. 

Key words: Burrow temperature, Energetics, Ornithorhynchus anatinus, Platypus, Tasmania 



INTRODUCTION 



MATERIALS AND METHODS 



The platypus, Ornithorhynchus anatinus, 
inhabits the lakes, rivers and streams of eastern 
Australia from the Cooktown area in the north to 
Tasmania in the south. Over much of its range, the 
animal is found in alpine and tableland areas where, 
especially in winter, air temperatues fall well below 
freezing and water temperatures approach 0°C (Grant 
1995). Grant (1983a) suggested that under such 
conditions, the microhabitat of platypus burrows is of 
great importance for energy conservation. Even in an 
unoccupied artificial burrow the insulation of layers 
of earth was found to provide significant buffering 
effect against outside ambient temperature changes 
both in winter and in summer (Grant 1976). (Grant 
1983b) suggested a further modifying effect of the 
animal's presence on the microhabitat temperature, 
elevating it several degrees above that of an unoccupied 
burrow. 

In this study ambient temperatures in 
occupied platypus burrows at a sub-alpine Tasmanian 
lake were investigated. The use of time-depth recorders 
with integrated temperature sensors made it possible 
to determine burrow temperatures during naturally 
occuring resting periods of the equipped animals. 



Field experiments were carried out at Lake 
Lea (41°30' S, 146°50' E), a sub-alpine lake in 
northwestern Tasmania. Information on burrow 
temperatures was obtained from nine individual 
platypuses (4 adult males, mass: 2.27 kg ± 0.26 (SD), 
5 adult females, mass 1.48 kg ± 0.07 (SD)) between 
November 1998 and January 2000. Platypuses were 
captured and processed following the methods outlined 
in Otley et al. (2000) and Bethge et al. (2003). 
Individuals were equipped with combined data logger- 
transmitter packages (max 62 mm x 28 mm x 18 mm, 
weight 50 g, Fig. 1) consisting of a specially designed 
standard transmitter (Faunatech, Eltham, Victoria) and 
a time-depth recorder (LTD 10, Lotek Inc., Canada). 
The packages were attached with glue (5 min-Araldite, 
Selleys Inc., Australia) to the guard fur of the lower 
back of the animals, just above the tail, following the 
method outlined in Serena (1994). Animals were 
released at the site of capture. After approximately two 
weeks the animals were relocated by radiotracking and 
recaptured on emergence and the devices were 
removed. 

The data loggers allowed measurement of 
ambient temperature in the range from 2 to 25°C with 



PLATYPUS BURROW TEMPERATURES 




Tbur 



Tw 
Ta 



15/6 16/6 17/6 18/6 19/6 20/6 21/6 22/6 23/6 24/6 25/6 26/6 27/6 



Figure 1: Winter sample data of water (Tw), air (Ta) and burrow temperatures (Tbur, derived from a 
time-depth recorder with integrated temperature sensor fitted to the back of a platypus; the temperature 
is only shown at times when the animal was in the burrow). 



an accuracy of 0.06°C. The devices were calibrated 
by the manufacturer (Equipment for temperature- 
calibration: Neslab RTE-2000 Bath/Circulator and 
Omega HH40 Thermistor/Thermometer). Temperature 
sensors were located at the back end of the devices 
and were facing backwards when the devices were 
fixed on the platypus's lower backs. Ambient 
temperature was measured in two-minute intervals for 
11 days each. While foraging, the sensors measured 
water temperatures. In resting platypuses, ambient 
temperatures close to the animals' bodies (approx. 5 
mm from above the fur) were recorded. The resting 
period was defined as the time span between the end 
of the last dive of a foraging trip (detected by the depth 
sensor of the time-depth recorders) and the beginning 
of the first dive of the following foraging trip. Burrow 
temperatures, i.e. ambient temperatures during resting 
periods, were recorded in two minute intervals for a 
total of 61 resting periods and were averaged over the 
period of resting. Resting periods ranged from 5.45 to 
27.20 hours. 

All investigated platypuses occupied burrows 
in consolidated steep or gently sloping earth banks of 
the lake or along associated creeks. Water and air 
temperatures at Lake Lea were recorded in two-hour 



intervals using archival tags (HOBO Thermocouple 
logger and Stowaway Temperature Logger, Onset 
Computer Corp., USA). Water temperature was 
measured in the lake while air temperature was taken 
in a wind shaded forest patch nearby. 



RESULTS 

Mean burrow temperatures were 17.5 and 
14.2°C (SD=2.76 and 0.89, respectively, n=9) in 
summer and winter, respectively, and ranged between 
12.2 and 22.8°C for individual resting periods. In 
winter, burrow temperature was held fairly constant 
over the resting period while in summer larger 
variations were observed. A low but significant 
correlation between air temperature and burrow 
temperature was found with higher air temperatures 
resulting in higher burrow temperatures (p=0.003, 
n=6 1 ). Ambient air temperatures ranged between -4°C 
and 31°C and water temperatures between 0°C and 
29°C depending on season. Samples of measured 
burrow temperatures and corresponding air and water 
temperatures are shown in Fig. 1 and Fig. 2 for winter 
and summer, respectively. 



274 



Proc. Linn. Soc. N.S.W., 125, 2004 



P. BETHGE, S. MUNKS, H. OTLEY AND S. NICOL 



Tbur 




_, , , , , , , , , , r 

7/1 8/1 9/1 10/1 11/1 12/1 13/1 14/1 15/1 16/1 17/1 18/1 19/1 20/1 



Date 



Figure 2: Summer sample data of water (Tw), air (Ta) and burrow temperatures (Tbur, derived from a 
time-depth recorder with integrated temperature sensor fitted to the back of a platypus; the temperature 
is only shown at times when the animal was in the burrow). 



DISCUSSION 

Grant (1983b) suggested that platypus 
burrows act as microenvironments, buffering the 
animals against the rigours of below-freezing air 
temperatures in winter, and modifying the effects of 
high summer temperatures. Accordingly, we found that 
in winter, burrow temperatures at Lake Lea were up 
to 18°C higher than outside air temperatures (Fig. 1). 
In summer, the burrows at Lake Lea clearly buffered 
high midday temperatures of over 25°C (Fig. 2). These 
findings are in line with results by Grant (1976) and 
Munks (personal communication). In winter, 
unoccupied artificial burrow temperatures in the upper 
Shoalhaven River, NSW, averaged around 14°C (this 
study: 14.2°C) despite the fact that ambient air 
temperatures dropped as low as -5°C. During summer 
the temperature of an unoccupied artificial burrow 
averaged around 18°C (this study: 17.5°C) with air 
and water temperatures being several degrees higher 
(Grant 1976, Grant 1995). Munks (personal 



communication), while monitoring the burrow of a 
breeding platypus in lowland Tasmania, recorded a 
mean burrow temperature of 16.5°C (range 12.5 to 
20°C) during late summer/early autumn. 

The consistency of these data from different 
sites suggests that platypus burrow temperatures are 
fairly constant regardless of habitat. Whether this is a 
consequence of the metabolic heat produced by the 
animals or mainly of physical characteristics of their 
burrows, remains unclear. Results of Grant (1976) from 
unoccupied burrows are in line with findings presented 
here from occupied burrows. This suggests that - at 
least in burrows located in consolidated earth banks - 
physical characteristics of the burrow are more 
important for burrow temperature than the absence or 
presence of the animal. This view is supported by the 
significant correlation between air temperature and 
burrow temperature found in this study. 

However, Munks (personal communication) 
reported peak burrow temperatures when the mother 
returned to the nest to suckle her young. Also, Grant 
(1983b) suggested that the animal's presence further 



Proc. Linn. Soc. N.S.W., 125, 2004 



275 



PLATYPUS BURROW TEMPERATURES 



elevates the microhabitat temperature of the burrow. 
In captivity, Grant (1976) observed, that the 
temperature in an uninsulated plywood nest box rose 
around 1 to 2°C above ambient temperature when an 
animal was inside. 

We suggest that these different observation 
are a consequence of different burrow characteristics. 
In this study, all investigated platypuses occupied 
burrows in consolidated earth banks. Under such 
conditions, the insulation properties of the surrounding 
earth and of the nesting material inside the burrow are 
most likely the main factors determining burrow 
temperature. A fairly constant burrow temperature may 
of course be more critical during the breeding period 
(Grant, personal communication), which makes deep 
earth burrows ideal for nesting. 

A different situation, however, might occur 
in burrows, which are closer to the surface or above 
ground. Otley et al. (2000) reported that 25 % of 
burrows at Lake Lea were located within dense 
vegetation, such as sphagnum and button grass. The 
insulation properties of such burrows would be 
expected to be poor compared to underground earth 
burrows. How animals cope with high thermal stress 
in vegetation burrows and if they use this sort of burrow 
site regardless of season or even during nesting requires 
further investigation. 



ACKNOWLEDGMENTS 

This work was supported by the Australian 
Research Council, an Overseas Postgraduate Research 
scholarship by the University of Tasmania and a doctoral 
scholarship by the DAAD (Deutscher Akademischer 
Austauschdienst, Germany ,Hochschulsonderprogramm III 
von Bund und Landern'). The field work was carried out 
under permit from the Department of Parks, Wildlife and 
Heritage, Tasmania, the Inland Fisheries Commission, 
Tasmania and the University of Tasmania Ethics Committee. 
Thanks to all those who assisted with the field work and to 
Mr H. Burrows for access to private land. 



REFERENCES 

Bethge, P., Munks, S., Otley, H.and Nicol, S. (2003) 

Diving behaviour, dive cycles and aerobic dive 
limit in the platypus Ornithorhynchus anatinus. 
Comparative Biochemistry and Physiology A 
136/4, 799-809. 

Grant, T.R. (1976). Thermoregulation in the Platypus, 
Ornithorhynchus anatinus. PhD thesis, 
University of New South Wales, Australia. 

Grant, T.R. (1983a). Body temperatures of free-ranging 
platypuses, Ornithorhynchus anatinus 
(Monotremata), with observations on their use 
of burrows. Australian Journal of Zoology 31, 
117-122. 

Grant, T.R. (1983b). The behavioural Ecology of 

Monotremes. In , Advances in the Study of 
Mammalian Behaviour' (Eds J.F. Eisenberg and 
D.G. Klieman). The American Society of 
Mammalogists, Special Publication Vol. 7, pp 
360-394. 

Grant, T.R. (1995). The platypus. A unique mammal. 

University of New South Wales Press, Sydney. 

Otley, H.M., Munks, S.A. and Hindell, M.A. (2000). 
Activity pattern, movements and burrows of 
platypuses {Ornithorhynchus anatinus) in a sub- 
alpine Tasmanian lake. Australian Journal of 
Zoology 48, 701-713. 

Serena, M. (1994). Use of time and space by platypus 
{Ornithorhynchus anatinus: Monotremata) 
along a Victorian stream. Journal of Zoology 
232, 117-130. 



276 



Proc. Linn. Soc. N.S.W., 125, 2004 



Ultrasonography of the Reproductive Tract of the Short-beaked 
Echidna (Tachyglossus aculeatus) 

D. P. HlGGINS 

Faculty of Veterinary Science, B01, University of Sydney NSW 2006 
(damienh@vetp.usyd.edu.au) 



Higgins, D.P. (2004). Ultrasonography of the reproductive tract of the short-beaked echidna 
{Tachyglossus aculeatus). Proceedings of the Linnean Society of New South Wales 125, 277-278. 

We describe a brief investigation of ultrasonography as a tool to monitor reproductive activity and to 
determine the sex of short- beaked echidnas {Tachyglossus aculeatus). We found trans-abdominal ultrasound 
to be of limited use for monitoring ovum development but it appears to be useful for imaging the uterus. We 
also found ultrasonography to be a useful tool to confirm the sex of echidnas by visualizing the testis. 

Manuscript received 18 August 2003, accepted for publication 8 January 2004. 

KEYWORDS: abdomen, echidna, monotreme, reproduction, Tachyglossus, testis, ultrasound, uterus. 



Here we describe a brief investigation of 
ultrasonography as a tool to monitor reproductive 
activity and to determine the sex of short- beaked 
echidnas {Tachyglossus aculeatus). Griffiths (1968) 
described the gross anatomy of the reproductive tract 
of the female echidna. An ovum of 3-4 mm diameter 
is ovulated from one of the two flat, sauropsid-like 
ovaries, which lie ventrocaudal to the kidneys 
(Griffiths 1968). Although only one ovum is ovulated 
in the echidna, Flynn (1930) reported that up to three 
large ova and several much smaller ova may occur on 
the ovary. Hughes and Carrick (1978) concluded from 
Hill and Gatenby (1926), Caldwell (1887) and Flynn 
and Hill (1939) that the ovum has a vitelline membrane, 
a zona pellucida and a proalbumen which may be 
analogous to the liquor folliculi of the graafian follicle, 
but has no follicular antrum. During its passage down 
the fallopian tube, the ovum swells to 5 mm diameter. 
The shell membrane is first laid down in the fallopian 
tube and later thickens in the uterus. The egg absorbs 
fluid in-utero and expands from 6.5mm diameter to 
15 mm x 13 mm. 

Ten short- beaked echidna carcasses were 
placed in dorsal recumbency. A portable ultrasound 
machine with a 7.5 Mhz linear transducer (SSD- 500, 
Aloka, Japan) was used to image the abdomen. Results 
were confirmed by dissection. An additional nine 
echidnas were then anaesthetized and examined in a 
similar fashion. Positioning the transducer on the 
ventral abdominal wall, lateral to the epipubic bones 
avoided the need to shave the hair of the pseudopouch 
and minimised interference by intestinal gas. 



Dissection confirmed that the gonads lie against the 
dorsal body wall, dorsal to the cranial ends of the 
epipubic bones. Ovaries of freshly dead echidnas 
lacked grossly visible developing ova. Of frozen and 
thawed bodies, which generally had poorer tissue 
contrast, ovaries and ova were not visible by 
ultrasonography. A structure in the expected location 
of the ovaries and comprising several 2- 3 mm 
diameter, thin walled, echolucent bodies was 
sometimes visible in living echidnas during the 
breeding season, however, the scarcity of surrounding 
interstitium made repeatable identification of 
individual putative ova very difficult. In addition, the 
small intestine frequently cast gas shadows over the 
gonads, reducing their visibility. It is likely that trans- 
rectal ultrasound would improve visualization of the 
ovaries but may be of limited use in serial observations, 
where the extent of manual or chemical restraint 
required may introduce variations to the reproductive 
cycle (Clarke and Doughton 1983, River and Rivest 
1991). The entire oviducts of reproductively active live 
and dead animals were clearly visible, especially when 
adjacent to a full bladder. Ova were not seen in the 
oviducts of any of our animals. Testes appeared as 15 
to 25mm long, ovoid, homogenous, soft tissue 
structures and, when present, were always visible 
caudal to the kidney and, on the left side, dorso-caudal 
to the mobile, spherical portion of the spleen. Due to 
their similar appearance on ultrasound, both the spleen 
and testis were sighted on the left before the left testis 
was identified. 



ULTRASONOGRAPHY IN ECHIDNA REPRODUCTIVE STUDIES 



In conclusion, we found trans-abdominal 
ultrasound to be of limited use for monitoring ovum 
development but it appears to be useful for imaging 
the uterus. In sexing echidnas, the inability to extrude 
or palpate a phallus does not confirm its absence, and 
other characteristics such as absence of a pseudo-pouch 
or presence of spurs may not be reliable indicators of 
sex, therefore the gender of echidnas in captive 
collections is sometimes mistaken. We found 
ultrasonography to be a useful tool to confirm the sex 
of echidnas in these circumstances. 



ACKNOWLEDGMENTS 

We thank Dr Marianne Offner for her assistance 
in interpretation of ultrasonographs, Medtel for the provision 
of the Aloka SSD-500 ultrasonography machine, and the 
Zoological Parks Board of NSW, Novartis Australia and the 
Winifred Scott Foundation for their financial support. 



REFERENCES 

Caldwell, W. H. (1887) The embryology of Monotremata 
and Marsupialia- Parti. Philosophical 
Transcripts of the Royal Society. 178(B), 463- 
480. 

Clarke, I. J. and Doughton, B. W. (1983) Effect of various 
anaesthetics on the resting plasma 
concentrations of lutienising hormone, follicle 
stimulating hormone and prolactin in 
ovariectomised ewes. Journal of Endocrinology. 
98, 79-89. 

Flynn, T. T. (1930). On the unsegmented ovum of the 
echidna (Tachyglossus) Quarterly Journal of 
Microscopical Science. 74, 119-131. 

Flynn, T. T. and Hill, J. P. (1939) The development of the 
Monotremata Part VI -Growth of the ovarian 
ovum, maturation, fertilisation and early 
cleavage. Philosophical Transcripts of the Royal 
Society (London) XXIV (6), 571-578. 

Griffiths, M. E. (1968) 'The Echidna.' (Pergamon Press: 
UK). 

Hill, J. P. and Gatenby, J. B. (1926). The corpus luteum of 
the Monotremata. Philosophical Transcripts of 
the Royal Society (London) II, 7 1 5-762. 

Hughes, R. L. and Carrick, F. N. (1978). Reproduction in 
female monotremes. Australian Zoologist 20, 
233-254. 

River, C. and Rivest, S. (1991). Review Article. Effect of 
stress on the activity of the hypothalamic- 
pituitary- gonadal axis: peripheral and central 
mechanisms. Biology of Reproduction 45, 523- 
532. 



278 Proc. Linn. Soc. N.S.W., 125, 2004 



Excretion Profiles of Some Reproductive Steroids in the Faeces 

of Captive Female Short-beaked Echidna (Tachyglossus 

aculeatus) And Long-beaked Echidna (Zaglossus sp.) 

D.P. Higgins, G. Tobias, G.M. Stone 
Faculty of Veterinary Science, B01, University of Sydney NSW 2006 (damienh@vetp.usyd.edu.au) 



Higgins, D.P., Tobias, G. and Stone, G.M. (2004). Excretion profiles of some reproductive steroids in the 
faeces of captive female short-beaked echidna {Tachyglossus aculeatus) and long-beaked echidna 
(Zaglossus sp.). Proceedings of the Linnean Society of NSW 125, 279-286. 

We evaluated and applied an existing faecal reproductive steroid extraction and radio-immunoassay (RIA) 
procedure to samples from captive short-beaked (Tachyglossus aculeatus) and long- beaked (Zaglossus sp.) 
echidnas. Steroids were extracted from faeces with diethyl ether, resuspended in 80% methanol and lipids 
removed with petroleum ether. The methanol fraction was purified and assayed for progestins or oestrogens, 
results corrected for procedural losses and converted to ng/ g dry weight of faeces. One T. aculeatus was 
injected with radiolabelled and natural progesterone and faecal extracts were subjected to high- performance 
liquid chromatography (HPLC) to allow partial identification of radiolabelled and RIA- reactive metabolites. 
The major RIA-reactive substance and the major labelled [ 14 C] compound co-eluted with progesterone. An 
additional RIA-weak compound co-eluted with 20[3-dihydroxyprogesterone, and three additional RIA-weak, 
radio-labelled compounds eluted but were not identified. Increases in faecal progestin of echidnas occurred 
at 17 ± 3 (n = 5), 33 ± 3 (n = 4) and 48 (n = 1) day intervals, supporting a cycle length of approximately 17 
or 33 days. However, further study incorporating more animals, behavioral observations and more frequent 
sampling of faecal oestrogens is required to produce more definitive results. 

Manuscript received, 18 August 2003, accepted for publication 8 January 2004. 

KEYWORDS: Faecal reproductive steroids, HPLC, monotreme, oestrogen, progestin, 
radioimmunoassay, Tachyglossus aculeatus, Zaglossus. 

INTRODUCTION gestational length, as in pygmy possums (Cercartetus 

spp), brown antichinus (Antichinus stuartii), eastern 

The short- beaked echidna (Tachyglossus quolls (Dasyuris viverrinus) (Tyndale- Biscoe 1973) 

aculeatus) is widespread within Australia and New and bent-winged bats (Miniopterus spp) (Wimsatt 

Guinea. The long- beaked echidna (Zaglossus bruijnii) 1 969) . 

is restricted to the highlands of New Guinea where it Longitudinal studies better define 

is endangered by human interference (Flannery 1990). reproductive cycles and illustrate inter-individual 

Despite more than 100 years of captive husbandry, it variation than cross-sectional studies, which are more 

is rare for these animals to breed in captivity (Augee conveniently applied to wild animals (Lasley 1985). 

et al. 1978; Boisvert and Grisham 1988) and However, even captive echidnas are cryptic and curl 

knowledge of the timing and hormonal control of the into a tight ball when threatened, making difficult the 

reproductive cycles of monotremes is limited. The frequent collection of blood, urine or urogenital swabs 

presence of a luteal phase is generally accepted, based without anaesthesia or forceful restraint, which may 

on histological evidence (Hill and Gatenby 1926; cause variation of reproductive cycles and behaviour 

Griffiths 1968; Hughes and Carrick 1978; Griffiths (Clarke and Doughton 1983; Rivier and Rivest 1991; 

1984) but its role and duration is unknown. In addition, Cleva et al 1994). Non- invasive faecal reproductive 

observations of gestation range from greater than 10 steroid assays have been used to describe the 

days (Carrick 1977) to 28 days (Broom 1895) after reproductive cycles of many species. This paper reports 

mating. Griffiths (1984) speculated that, like some the initial assessment and application of a faecal 

reptiles and bats (Racey and Potts 1970), female reproductive steroid assay as a non- invasive technique 

echidnas may store sperm, or that torpor may alter for the first sequential study of female echidna 



FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS 



reproductive endocrinology. 



MATERIALS AND METHODS 

Animals and housing 

Study animals were six female T. aculeatus, 
aged between 4 and 7 years and of 3 to 5 kg 
bodyweight, and three female Zaglossus bruijnii 
(probably Z. bartoni of Flannery and Groves 1998), 
aged between 20 and 32 years and of 6.5 to 14 kg 
bodyweight, all from the Taronga Zoo collection. The 
study was conducted in two phases: From May to 
September 1995, two T. aculeatus and all Zaglossus 
sp. were housed in two indoor enclosures with reverse 
cycle seasonal lighting and in the continual presence 
of a male of their species. From June to October 1997, 
four female T. aculeatus were housed outdoors, in 
adjacent 5.1 x 6.3 m enclosures. These females were 
housed individually to accommodate the solitary nature 
of the animal (Augee et al. 1975; Abensperg-Traun 
1991) and to facilitate identification of the source of 
faeces. A male T. aculeatus had access to all four 
enclosures through magnetically controlled doors until 
they failed, after which he was manually rotated, daily, 
between enclosures. Sixty-centimeter deep woodchip 
substrate, half pipes, tables, tree branches and logs were 
provided as shelter. 

Feeding and sample collection 

Animals were fed daily slurry of minced beef, 
egg, cereal, vitamin and mineral supplements and 
sufficient unprocessed bran to produce firm stools. 
Initially, 1mm x 1mm x 3mm food grade polyethylene 
pellets (Hoechst Industries, Australia) were added to 
the food of the indoor groups to identify the source of 
faeces. It appeared possible that not all pellets were 
being excreted, therefore the feeding of pellets was 
discontinued and for the course of the study each 
animal from the indoor groups was placed in a separate 
room for 24- 48 h once weekly and faeces were 
collected. Blue food dye (8 mg/day, Hexacol Brilliant 
Blue FCF Supra 75328, Pointing Hodgsons Pty Ltd, 
Australia), was added to the food of the outdoor female 
animals to make faeces more visible, and all visible 
faeces were collected daily. Samples were handled 
using latex gloves and stored in plastic zip- lock bags 
at -20°C for up to one year. 

Extraction and purification 

Due to the need to separate echidnas for 
sample collection in the first phase of the study, the 
sampling interval for progestin excretion profiles of 
echidnas housed indoors was 5 to 7 days. Sampling 



interval for progestin and oestrogen excretion profiles 
of echidnas housed individually outdoors in the second 
phase of the study was 1 to 3 days. The steroid 
extraction technique was based on a procedure used 
by Hindle and Hodges (1990). Each stool was finely 
chopped and mixed, then duplicate 0.5g samples were 
transferred to new glass vials (Econo Glas Vial, 
Packard, USA). Pieces were broken up using a glass 
rod, 5 ml diethyl ether (APS, Ajax Finechem, 
Australia) was added and vials were rotated for 30 min 
then centrifuged at 1500 G for 15 min at 4°C. The 
faecal sediment and aqueous portion were frozen in 
liquid nitrogen. Supernatant was decanted, evaporated 
at 30°C under nitrogen gas and reconstituted in 5 ml 
80%(v/v) methanol (80% MeOH) by rotation for 30 
min. Solutes were partitioned by addition of 5ml of 
petroleum ether (B.P. 40°C to 60°C, APS, Ajax 
Finechem, Australia), rotation for 20 min, and 
centrifugation at 1500 G for 15 min. The 80% MeOH 
fraction was aspirated and then stored at -20°C. 
Following extraction, faecal residue was dried at 100°C 
for 4 hours and weighed to determine dry matter 
content. 

Aliquots of 500 |il faecal extract in 80% 
MeOH were dried at 80°C under vacuum, reconstituted 
in 1ml 10% MeOH by agitation at 30°C for 30 min, 
and purified using Sep-Pak C18 Cartridges (Waters 
Scientific, Milford, USA) according to manufacturers 
recommendations. Eluants of 25%, 50%, 75%, 90% 
and 100% MeOH were collected and stored at -20°C. 

Radioimmunoassay (RIA) 

Duplicate 200 u.1 aliquots of eluates 
(unknowns) were dried and reconstituted in 200|ll 10% 
MeOH in IP buffer (0.03 1M Na 2 HP0 4 , 0.0 19M 
NaH 2 P0 4 .2H 2 0, 0.154M NaCl and 0.1%w/v gelatin, 
pH 7.4) by agitation at 30°C for 60 min. Radiolabeled 
steroids ([1,2,6,7- 3 H] progesterone in toluene ([ 3 H]P, 
96 Ci/mmol; Amersham Australia, Sydney, NSW) or 
[2,4,6,7- 3 H] oestradiol in toluene ([ 3 H]E, 104 Ci/mmol; 
Amersham Australia, Sydney NSW)) were dried and 
reconstituted in IP buffer to approximately 15000 dpm/ 
100 u.1. Our ovine antiserum to progesterone- 11a- 
hemisuccinate-BSA conjugate (1:55000 final dilution, 
#C-9817 Sirosera™, CSIRO Bioquest, Blacktown, 
Australia), cross-reacted with progesterone (100%), 
11(3- hydroxyprogesterone (32.5%), corticosterone 
(18.8%), 20a- hydroxy-4-pregnane-3-one (0.7%), 
17a- hydroxyprogesterone (0.2%), 20(3- hydroxy-4- 
pregnane-3-one (0.2%), pregnenolone (0.2%), 
oestradiol (<0.2%), testosterone (<0.2%), Cortisol 
(<0.2%) (Curlewis, Axelson and Stone, 1985). Our 
ovine antiserum to 17(3- oestradiol-6- 
carboxymethyloxime-BSA (1:100000, #9757 



280 



Proc. Linn. Soc. N.S.W., 125, 2004 



D.P. HIGGINS, G. TOBIAS AND G.M. STONE 



Sirosera™, CSIRO Bioquest, Blacktown, Australia), 
cross-reacted with oestradiol (100%), oestrone 
(10.8%), oestriol (2.3%), oestradiol- 17a (<0.1%), 
progesterone (<0.1%), testosterone (<0.1%), 
androstenedione (<0.1%), Cortisol (<0.1%), 
corticosterone (<0.1%) (Curlewis 1983). Standards 
were generated from two overlapping doubling 
dilutions of progesterone (BDH Chemicals, Australia) 
from 500 - 7.84 pg/100 jn.1 or oestradiol (BDH 
Chemicals, Australia) from 500 - 1.96 pg/100 ul. 

Reactions contained 100 ul radiolabeled 
steroid, 100 ul antiserum and either 200 ul of unknown 
in 10% MeOH in IP buffer or 100 ul 20% MeOH in 
IP buffer and 100 ul of standard. The resulting 400 ul 
was vortexed for 30 sec and incubated at 4°C for 18 h. 
Triplicate "total" (200 ul IP buffer, 100 ul 
radiolabeled hormone in IP buffer, 100 ul 20% MeOH 
in IP buffer), "non-specific binding" (200 ul IP buffer, 
100 ul radiolabeled hormone in IP buffer, 100 ul 20% 
MeOH in IP buffer) and "B " (100 ul IP buffer, 100 
Ul antiserum in IP buffer, 100 ul radiolabeled 
hormone in IP buffer, 100 ul 20% MeOH in IP buffer) 
standards were processed simultaneously with 
unknowns and standards. 

Free radiolabeled hormone was removed 
from all except "total" solutions by incubation for 10 
min at 4°C with 500 ul of charcoal/ dextran solution 
(0.25% w/v Norit-A filtered activated charcoal powder, 
Matheson, Coleman and Bell, USA) and 0.025% w/v 
dextran T70 (Pharmacia Fine Chemicals, Sweden) 
suspended in IP buffer). In place of the charcoal/ 
dextran solution, 500 ul of milli Q water was added to 
"total" solutions. After centrifugation at 1500 G for 
10 min at 4°C, the supernatant was decanted and its 
radioactivity measured as counts per minute (cpm) on 
a Beckmann LS 6500 Liquid Scintillation Spectrometer 
(Beckmann Instuments Inc, CA, USA.), which then 
converted cpm to disintegrations per minute (dpm) 
using an external standard. 

High- performance liquid chromatography (HPLC) 
of excreted metabolites 

One female T. aculeatus was injected intra- 
peritoneally with 5 mCi of [4- 14 C] progesterone ([ 14 C]P, 
48.9 mCi/mmol, NEN Dupont, USA) and 2 mg of 
natural progesterone in 30% (v/v) propylene glycol in 
isotonic saline. Eight 0.5 g faecal samples were 
obtained two days after injection. Extracts from these 
samples were pooled into two samples and subjected 
to sep-pak chromatography. Eluates of 2252 dpm and 
2440 dpm were dried under N 2 gas, reconstituted in 
75% acetonitrile, filtered and subjected to HPLC 
(K65B HPLC system, ETP Kortec, Australia) at a flow 
rate of 0.5 ml per minute, using 61% acetonitrile at a 



pressure of 2250 psi at room temperature. Fractions 
were collected every 30 sec for 22 min, then every 
minute for 19 min, then every 10 min for 20 min. 
Absorbance at 240nm was measured, to monitor the 
separation of steroids with a 4-ene-3-ketone structure. 
Elution time of progesterone was identified using [ 3 H]P 
and a progesterone standard and the column was 
calibrated for testosterone, androstenedione, 
progesterone and 20a dihydroprogesterone. 

Assessment of extraction, purification and RIA 
procedures 

Three different solvents were tested for use 
in the extraction process. Faeces containing metabolites 
of injected radiolabeled and natural progesterone were 
agitated in 90% MeOH, 80% MeOH or diethyl ether, 
and partitioned with petroleum ether as described 
above. The three solvents and their petroleum ether 
portions were assayed for progestins as above. 

The Sep-pak chromatography elution profile 
for oestrogen calculated by Spanner et al. (1997) was 
assumed for this study. The elution profile for progestin 
was determined by Sep-pak chromatography of 
solutions containing 200 fmol [ 3 H]P, using the series 
of MeOH dilutions described previously or the same 
series of dilutions of ethanol (EtOH). Co-elution of 
metabolites of faecal origin with progesterone was 
determined by adding 25000 dpm [ 3 H]P to duplicate 
faecal extracts from five female T. aculeatus and 
subjecting these to Sep-pak chromatography. 

Sample steroid recovery was estimated by 
adding 25000 dpm [ 3 H]P or 30000 dpm [ 3 H]E to 
respective samples and then performing the extraction. 
Duplicate 50 ul aliquots of purified 80% MeOH extract 
were combined with 500 ul Milli-Q water and 5 ml of 
scintillation fluid. Triplicate "total" vials were 
prepared, each containing 100 ul of radiolabeled 
hormone solution, 50 ul 80% MeOH, 400 ul Milli-Q 
water and 5 ml of scintillation fluid. Triplicate "blank" 
vials were prepared, each containing 50 ul 80% MeOH, 
500 ul Milli-Q water and 5 ml of scintillation fluid. 
Radioactivity was measured and percentage recovery 
was calculated by the formula: 

R = [400(d-B)/ (T-B)] x 100 

where R = percentage recovery (%), d = sample dpm, 
B = mean "blank" dpm, T = mean "total" dpm. 

To assess parallelism, faecal extracts from 
three faecal samples were reconstituted and diluted 
twofold and fourfold in 10% MeOH in IP buffer. 
Standards were similarly diluted and all dilutions were 
assayed for progestins as described above. Spanner et 
al. (1997) estimated parallelism of the oestradiol assay. 



Proc. Linn. Soc. N.S.W., 125, 2004 



281 



FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS 



The inter- assay coefficients of variation for 
progesterone and oestradiol assays were taken as the 
mean of the coefficients of variation of repeated (n=2), 
duplicated extraction and assay of 6 and 4 randomly 
chosen samples, respectively. 

The intra- assay coefficient of variation was 
estimated from the mean of two coefficients of 
variation, each calculated from five concurrent 
replicate extractions and assays of two randomly 
chosen samples. The intra-assay coefficient of variation 
was estimated for two progesterone extraction methods 
to determine the homogeneity of steroid in the stool. 
In the first (unmixed) method, 5 samples were taken 
from an intact stool. In the second (mixed) method, 
the stool was finely chopped and mixed and each of 
the 5 samples consisted of at least 5 randomly chosen 
pieces from the mix. Spanner et al. (1997) estimated 
the intra-assay coefficient of variation of the oestradiol 
assay. 

Data analysis 

Standard curve generation and conversion of 
dpm to pg hormone/ scintillation vial were calculated 
with "Assayzap" (Biosoft, Cambridge). All other 
calculations and graphs were made using "Excel 5.0" 
(Microsoft, USA). Mean steroid recovery was 
calculated from the first 60 samples in each assay. 
Mean recovery was used to correct results of progestin 
assays for procedural losses. As steroid recovery was 
more variable in oestrogen assays, results were 
corrected using a recovery value calculated for each 
individual sample. Faecal steroid peaks were defined 
as those values greater than 1.5 standard deviations 
from the mean of all values from that animal (Graham 
et al 1995). 



Assessment of progestin extraction, purification 
and RIA procedures 

As an initial solvent, ether extracted 37.0 ± 
4.3% (mean ± S.E.) more [ 14 C] labelled progestin than 
either 80% MeOH or 90% MeOH and was used in all 
subsequent extractions. Less than 10% of extracted 
steroid appeared in the petroleum ether fraction. Mean 
percentage recovery of [ 3 H]P through extraction and 
Sep-pak chromatography was 52% ± 7.17 (mean ± 
SD). Mean percentage recovery of [ 3 H]E was 30% ± 
13.7 (mean±SD). 

Almost all [ 3 H]P was recovered during Sep- 
pak chromatography. MeOH was chosen as the 
chromatography solvent as EtOH eluted the [ 3 H]P 
across a greater range of EtOH concentrations. Of 
recovered [ 3 H]P metabolites, 79.1% eluted in the 90% 
MeOH fraction and 93% eluted in the 75% MeOH 
and 90% MeOH fractions combined, with a mean 75% 
MeOH: 90% MeOH ratio (75:90 ratio) of 1:3.5. Of 
the progestin RIA- reactive faecal steroids recovered 
from the column, 85% was measured in the combined 
75% MeOH and 90% MeOH fractions with a mean 
75:90 ratio of 1:2.5. 

Correlation coefficients of parallelism curves 
for the progestin assay ranged from 0.988 to 1.000. 
Dose response curves for standards and extracts did 
not differ significantly (P>0.05) in slope. Sensitivity 
of the assay, as defined by 10% displacement from 
the Bo binding was 10 pg/assay tube. The intra- assay 
coefficients of variation were estimated to be 6.2% 
(mixed) and 25.7% (unmixed), therefore the mixed 
method was employed in all further extractions. The 
inter- assay coefficient of variation was estimated to 
be 14.9% for progesterone assays and 6.8% for 
oestradiol assays. 



RESULTS 

High pressure liquid chromatography 

The eluate with the highest RIA activity and 
moderate radioactivity was collected at 26 min, 
approximating the progesterone standard, which eluted 
at 25.5 min. [ 3 H]P eluted at 24 min. A moderately 
radioactive eluate with poor RIA activity that was 
collected at 22 min coincided with a 20(3- 
dihydroxyprogesterone standard, which has a low cross 
reactivity with the antiserum. Other [ i4 C]-labelled, 
moderately RIA-reactive compounds that eluted at 30, 
36 and 38 min and one [ 14 C]-labelled, weakly RIA- 
reactive compound that eluted at 22 min were not 
identified. One RIA-reactive compound that eluted at 
41 min did not co-elute with a [ l4 C]-labelled metabolite 
and this substance is yet to be identified. 



Faecal progestins 

Maximum and minimum faecal progestin 
concentrations from each of the six T. aculeatus ranged 
from 480 to 1 800 ng/g dry weight faeces (mean 860 
ng/g) and 5 to 100 ng/g dry weight faeces (mean 71 
ng/g), respectively. Intervals between samples that 
contained progestin peaks clustered at 17 ± 3 (n = 5), 
33 ± 3 {n = 4) and 48 (n = 1) days. Maximum and 
minimum faecal progestin concentrations from each 
of the three Zaglossus sp. ranged from 260 to 500 ng/ 
g dry weight faeces (mean 420 ng/g) and 10 to 70 ng/ 
g dry weight faeces (mean 40 ng/g), respectively. Two 
Zaglossus sp. produced two peaks each and the 
intervals between samples that contained these were 
28 and 70 days. The third produced one peak. 

Faecal Oestrogens 

Maximum and minimum faecal oestrogen 
concentrations from the four animals ranged from 21 



282 



Proc. Linn. Soc. N.S.W., 125, 2004 



D.P. fflGGINS, G. TOBIAS AND G.M. STONE 



to 45 ng/g dry weight faeces (mean 33 ng/g) and 3 to 
14 ng/g dry weight faeces (mean 7 ng/g), respectively. 
Intervals between adjacent oestrogen peaks were 8, 
19, 24 and 30 days apart. Fluctuations approaching 
1.5 SD above the mean were common and made 
difficult the detection of any possible cyclic activity 
as intervals between these ranged from 4 to 16 days. 

Combined profiles 

Of the 8 oestrogen peaks, 7 were associated 
with progestin increases to concentrations less than 
1.5 SD above the mean. Combined oestrogen and 
progestin profiles from two T. aculeatus are shown 
in Figures 1 and 2. 



DISCUSSION 

Feeding and sample collection 

Plastic pellets were less suitable as a faecal 
marker than the blue food dye. T. aculeatus consumed 
95% of pellets placed in their food while Zaglossus 



sp. consumed less than 50%. Not all ingested pellets 
were recovered, indicating that pellets were being 
retained or faeces were remaining undetected. Food 
containing the blue dye was readily eaten and faeces 
containing the dye were considerably more detectable 
than those without. Passage time of the plastic pellets 
ranged from 12 hours to greater than 48 hours. 

High pressure liquid chromatography 

The strong antiserum cross-reactivity of the 
substance which eluted at 26 min, and its proximity to 
the elution time of the progesterone standard, makes 
progesterone its likely identity, however further 
confirmation of this would be desirable as we are 
unable to explain the elution of [ 3 H]P 2 min earlier. 

The cross-reactive metabolites that were less 
polar than progesterone were not identified. These 
substances may contribute to the difference in 75:90 
ratio between the Sep-pak elution profiles of [ 3 H]P and 
progestins of faecal origin as the 75%MeOH, or less 
polar, component was less in the [ 3 H]P profile. As a 
priority, future studies should identify the RIA-reactive 




56 64 65 68 69 70 72 75 76 77 79 81 83 86 89 92 94 96 

day 



■■■ progestin 
—♦—oestrogen 



Figure 1. Combined faecal oestrogen and progestin profiles of one T. aculeatus over a 40-day period 
showing alternating progestin and oestrogen peaks greater than 1.5 SD above the mean, suggesting an 
oestrous cycle of 29 days. Also visible are increases less than 1.5 SD above the mean, suggesting concurrent 
progestin and oestrogen rises at 65, 79 and 94 days with interceding raised progestin/ lowered oestrogen 

periods surrounding 70 and 89 days, suggesting two cycles of 15 days. = mean faecal progestin/ 

oestrogen concentration; = mean faecal progestin/ oestrogen concentration + 1.5 SD. 



Proc. Linn. Soc. N.S.W., 125, 2004 



283 



FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS 



900 
800 
700 



■M progestins 
— ♦— oestrogens 




25 



27 33 36 42 45 52 58 61 62 64 66 69 71 72 74 76 78 80 82 84 86 88 90 92 94 

days 



Figure 2. Combined faecal oestrogen and progestin profiles of one T. aculeatus over a 67-day period 
showing alternating progestin and oestrogen peaks greater than 1.5 SD above their respective mean, 
suggesting an oestrous cycle of approximately 32 days. Also visible are additional fluctuations of oestrogen 
concentration at 52, 78, 82 and 86 days and progestin at 36 days, which hinder clear interpretation of 

oestrous cycles. = mean faecal progestin/ oestrogen concentration; = mean faecal 

progestin/ oestrogen concentration + 1.5 SD. 



compound that did not correspond to a radiolabeled 
metabolite. A similar examination of oestrogen 
metabolites would assist interpretation of oestradiol 
assays. 

Extraction and recovery 

As homogeneity of steroid in the faeces was 
poor, mixing of the stool before sampling was 
necessary to reduce intra-assay variance. Though 
diethyl ether extracted the most steroid, recovery 
through extraction was low and variable in this study, 
especially in oestradiol assays. We attempted to correct 
for this by correcting for procedural losses using 
individual recovery values for each sample in 
oestrogen assays but progestin assay data were 
corrected using a mean recovery value. Use of 
individual recovery values for progestin samples may 
have improved interpretation of data. 

MeOH concentrations exceeding 5% in the 
RIA incubations considerably reduced steroid- 
antibody binding. At each corresponding 



concentration, EtOH had a greater effect on steroid- 
antibody binding than MeOH. Reconstitution of eluates 
in 10% MeOH to produce a final concentration of 5% 
MeOH in the assay provided adequate steroid solubility 
and minimised interference with steroid-antibody 
binding. 

Progestin and oestradiol profiles 

Mean, maximum and minimum progesterone 
and oestrogen values varied among animals, indicating 
that this technique may be unsuitable for assessing the 
status of an animal from a single measurement. The 
small number of animals available for the study and 
the need for further work to identify an ti serum-reactive 
metabolites limits the conclusions that can be drawn 
from the sequential data obtained in this study. 
However, the lack of knowledge in this area makes 
some trends worthy of comment for consideration in 
future work. 

The intervals between subsequent progestin 
peaks in this study suggest a progesterone periodicity 



284 



Proc. Linn. Soc. N.S.W., 125, 2004 



D.P. fflGGINS, G. TOBIAS AND G.M. STONE 



of 16-17 days. However, as there was no clear pattern 
in oestrogen excretion, we could not determine whether 
this reflects concurrent vitelline progesterone and 
oestrogen peaks at 32- 34 day intervals with an 
interspersed luteal peak at 16-17 days (see figs 1 and 
2), or concurrent vitelline progesterone and oestrogen 
peaks at 16-17 day intervals with an interceding luteal 
phase with progestin increases below our arbitrary 
significance criterion. We expect that daily sampling 
and identification of potentially confounding RIA- 
reactive faecal steroids would be necessary to resolve 
this question. However, observations of fetal 
development add some support to the hypothesis of a 
17-day progesterone cycle. Decreasing blood 
progesterone is a precursor to parturition in many 
species of eutheria (Rowlands and Wier 1984) and 
metafheria (Tyndall-Biscoe and Renfree 1987), and 
to oviposition in many reptilia (Licht 1984). At 17 days 
the tammar wallaby fetus consists of 17- 20 somite 
pairs (Griffiths 1984), similar to the 19-20 somite pairs 
possessed by the echidna at oviposition (Hill and 
Gatenby 1926; Luckett 1976; Hughes and Carrick 
1978). Both young also exhibit similar stages of 
development at parturition or hatching 1 1 days later 
(Griffiths 1984). The similar rates of development in 
the last third of gestation and incubation suggests that 
the age of the echidna fetus at oviposition is 
approximately 17 days, consistent with a luteal phase 
of 16 to 17 days. 

The many irregular oestrogen fluctuations we 
measured could be inherent in the technique or 
indicative of follicular development and atresia. Hill 
and Gatenby (1926) described channels, from the 
vitellus to a well-developed lymphatic sinus in the 
ovarian medulla and histological features indicative 
of follicular regression in the platypus. 

The authors believe that this study provides 
a starting point for further work and suggest the further 
identification of progesterone and oestrogen 
metabolites and the comparison of faecal steroid 
concentrations with blood hormone concentrations, 
urogenital cytology, ultrasonography of the 
reproductive tract or behaviour in a controlled study 
accounting for the potential confounding effects of 
repeated physical or chemical restraint. 



ACKNOWLEDGEMENTS 

We thank the staff of the faculty of Veterinary 
Science, University of Sydney, in particular Michael Lensen, 
Irene van Ekris, Margaret Byrne and Geoff Dutton; and 
Taronga Zoo, in particular Margaret Hawkins, Kerry Foster, 
and Debbie Pritchard for their assistance. We also thank 
Michele Thums and the journal referees for their comments 



on the manuscript. The project would not have been possible 
without the help of the staff of Australian Mammals, Taronga 
Zoo and the Taronga Zoo Friends and without the financial 
assistance of Novartis, Australasia and the Winifred Scott 
Foundation. 



REFERENCES 

Abensperg-Traun, M. (1991) A study of home range, 

movements and shelter use in adult and juvenile 
echidnas {Tachyglossus aculeatus) in Western 
Australian wheatbelt reserves. Australian 
Mammalogy 14, 13-21. 

Augee, M.L., Bergin, T.J. and Morris, C. (1978). 

Observations on behaviour of echidnas at 
Taronga Zoo. Australian Zoologist 20, 121-129. 

Augee, M.L., Ealey, E.H.M. and Price, LP. (1975). 

Movements of echidnas Tachyglossus aculeatus 
determined by marking - recapture and radio 
tracking. Australian Wildlife Research 2, 93- 
101. 

Boisvert, M. and Grisham, J. (1988). Reproduction of the 
short beaked echidna at the Oklahoma City Zoo. 
International Zoo Yearbook 27, 103-108. 

Broom, R. (1895). Note on the period of gestation in 

echidna. Proceedings of the Linnean Society of 
NSW 10, 576-577. 

Carrick, F.N. (1977). Studies in the reproductive 

physiology of male marsupials. PhD thesis, 
University of New South Wales, Australia. 

Clarke, I.J. and Doughton, B.W. (1983). Effect of various 
anaesthetics on the resting plasma 
concentrations of lutienising hormone, follicle 
stimulating hormone and prolactin in 
ovariectomised ewes. Journal of Endocrinology 
98, 79-89. 

Cleva, G.M., Stone, G.M. and Dickens, R.K. (1994). 
Variation in reproductive parameters in the 
captive male koala. Reproduction, Fertility and 
Development 6, 713-719. 

Curlewis, J.D. (1983). Some interactions between gonadal 
steroid hormones and target organs in the male 
and female brush-tail possum (Trichosurus 
vulpecula). PhD thesis, University of Sydney, 
Australia. 

Curlewis, J.D., Axelson, M. and Stone, G.M. (1985). 
Identification of major steroids in the ovarian 
and adrenal venous plasma of the brush-tail 
possum (Trichosurus vulpecula) and changes in 
peripheral plasma levels of oestrogen and 
progesterone during the reproductive cycle. 
Journal of Endocrinology 105, 53-62. 

Flannery, T. (1990). 'Mammals of New Guinea.' (Robert 
Brown Associates: Australia). 

Flannery, T. F. and Groves, C. P. (1998). A revision of the 
genus Zaglossus (Monotremata, 
Tachyglossidae), with description of new 
species and subspecies. Mammalia. 62, 367- 
396. 



Proc. Linn. Soc. N.S.W., 125, 2004 



285 



FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS 



Graham, L.H., Goodrowe, K.L., Raeside, J.I. and Liptrap, 

R.M. (1995). Non-invasive monitoring of 

ovarian function in several felid species by 

measurement of fecal estradiol- 17b and 

progestins. Zoo Biology 14, 223-237. 
Griffiths, M.E. (1968). 'The Echidna' (Pergamon Press: 

UK). 
Griffiths, M. (1984). Mammals: Monotremes. In 

'Marshall's Physiology of Reproduction Vol 1, 

2nd ed' (Ed G.E. Lamming) pp. 351-385. 

(Churchill Livingstone: Edinburgh). 
Hill, J.P. and Gatenby, J.B. (1926). The corpus luteum of 

the Monotremata. Proceedings of the Zoological 

Society of London II, 715-762. 
Hindle, J.E. and Hodges, J.K. (1990). Metabolism of 

oestradiol-17a and progesterone in the white 

rhinoceros (Ceratotherium simum simum). 

Journal of Reproduction and Fertility 90, 571- 

580. 
Hughes, R.L. and Carrick, F.N. (1978). Reproduction in 

female monotremes. Australian Zoologist 20, 

233-254. 
Lasley, B.L. (1985). Methods for evaluating reproductive 

function in exotic species. Advances in 

Veterinary Science, Comparative Medicine 30, 

209-228. 
Licht, P. (1984). Reptiles. In 'Marshall's Physiology of 

Reproduction Vol 1 , 2nd ed' (Ed G.E. 

Lamming) pp. 206-282 (Churchill Livingstone: 

Edinburgh). 



Luckett, W.P. (1976). Fetal membranes of the 

Monotremata and the origin of mammalian 
viviparity. Anatomical Record 184, 466. 

Racey, P.A. and Potts, D.M. (1970). Relationship between 
stored spermatozoa and uterine epithelium in the 
pipistrelle bat (Pipistrellus pipistrellus) Journal 
of Reproduction and Fertility 22, 57-63. 

River, C. and Rivest, S. (1991). Review article. Effect of 
stress on the activity of the hypothalamic- 
pituitary- gonadal axis: peripheral and central 
mechanisms. Biology Reproduction 45, 523- 
532. 

Rowlands, I.W. and Weir, B.J. (1984). Mammals: non 

primate eutherians. In 'Marshall's Physiology of 
Reproduction Vol 1, 2nd ed' (Ed G.E. 
Lamming) pp. 455-658. (Churchill Livingstone: 
Edinburgh). 

Spanner, A., Stone, G. M.and Shultz, D. (1997). Excretion 
profiles of some reproductive steroids in the 
faeces of captive Nepalese red panda. 
Reproduction, Fertility and Development 9, 
565-570. 

Tyndale-Biscoe, H. (1973). 'Life of marsupials' (Edward 
Arnold: Australia). 

Tyndale-Biscoe, H. and Renfree, M. (1987). 

'Reproductive physiology of marsupials; 
monographs on marsupial biology.' (Cambridge 
University Press: Cambridge). 

Wimsatt, W.A. (1969). Some interrelations of 

reproduction and hibernation in mammals. 
Symposia of the Society for Experimental 
Biology 23, 511-549. 



286 



Proc. Linn. Soc. N.S.W., 125, 2004 



Anatomy of the Central Nervous System of the Australian 

Echidna 

M. Hassiotis 1 , G. Paxinos 2 and K.W.S. Ashwell 1 * 

'Department of Anatomy, School of Medical Sciences, The University of New South Wales, 2052, Sydney, 

NSW, Australia. 
2 Prince of Wales Medical Research Institute, The University of New South Wales, 2052, Sydney, NSW, 

Australia. 

*author to whom correspondence and proofs should be addressed. Postal address as above. 

Fax: 61 2 9385 8016, Phone: 61 2 9385 2482 

Email: k.ashwell@unsw.edu.au 



Hassiotis, M., Paxinos, G. and Ashwell, K.W.S. (2004). Anatomy of the central nervous system of the 
Australian echidna. Proceedings of the Linnean Society of New South wales 125, 287-300. 

Even from their gross appearance, the brain and spinal cord of the Australian echidna show unusual 
features. The spinal cord is one of the shortest ever recorded for any mammal, ending at the mid-thoracic 
level, a feature which may be related to the defensive posture of the echidna. The pattern of termination of 
unmyelinated afferents in the spinal cord as revealed by lectin labelling with the B4 isolectin from Griffonia 
simplicifolia is also quite different from that seen in placental mammals, with termination in patches within 
deeper layers of the dorsal horn. Within the brainstem, specializations of the trigeminal system are apparent 
with great enlargement of all trigeminal nuclei. The mesencephalic trigeminal nucleus also shows an unusual 
aggregation of neurons in a central midline position quite unlike therian mammals. While the dorsal thalamus 
of therian mammals shows compartmentation related to function , the dorsal thalamus of the echidna is 
remarkable for its lack of cytoarchitectural differentiation. Most of the high encephalization in this mammal 
is attributable to the highly gyrified cerebral cortex. This cortex is further distinguished by the positioning 
of the major functional areas (primary motor, somatosensory, visual and auditory areas) towards the caudal 
pole of the brain. 

Manuscript received 21 July 2003, accepted for publication 22 October 2003. 

KEYWORDS: cerebral cortex, echidna, monotreme, spinal cord, thalamus, trigeminal. 



INTRODUCTION (short-beaked echidna and platypus) (Iggo et al. 1 985 ; 

Scheich et al. 1986; Gregory et al. 1987, 1988, 1989; 
In this paper we will be reviewing what is Proske et al. 1998). This sensory modality utilizes the 
known about the anatomy of the central nervous system trigeminal system in both monotremes studied, 
of the Australian echidna (Tachyglossus aculeatus), To date, physiological and anatomical studies 

with special reference to those features with functional of peripheral sensory systems in this animal have 
relevance. Even at the level of gross inspection, the concentrated on peripheral receptors of the trigeminal 
central nervous system of the echidna is remarkable system. The short-beaked echidna is known to use its 
for the large size of the brain and the short relative sensitive snout as its major sensory tool. Anatomical 
length of the spinal cord. studies of this snout have revealed a rich distribution 

of unusual receptors on the tip (Andres et al. 1991; 

Manger and Hughes 1992). One of these, the gland 

PERIPHERAL RECEPTORS AND duct receptor system (Andres et al. 1991) or mucous 

ELECTRORECEPTION sensory gland (Manger and Hughes 1992), is present 

in both platypus and echidna and is thought to be 
One of the most remarkable features of involved in electroreception (Iggo et al. 1985; Scheich 
monotreme neurobiology, and one which touches on et al. 1986; Gregory et al. 1987, 1988, 1989). Despite 
trigeminal nuclei development and cortical this attention to snout receptors, very little attention 
organization, is the reported presence of has been given to the structure or function of central 
electroreception in two members of this subclass trigeminal pathways in any monotreme. 



ECHIDNA CENTRAL NERVOUS SYSTEM 



SPINAL CORD ANATOMY 

The spinal cord of the Australian echidna was 
examined by Ashwell and Zhang (1997). Even at the 
gross level, the spinal cord is notable because of its 
relatively short length, terminating at the level of the 
seventh thoracic vertebra (Figure la)(cf human spinal 
cord which terminates at the intervertebral disc 
between lumbar vertebrae 1 and 2). This may represent 



an adaptation to the pronounced vertebral flexure, 
which this mammal achieves when it adopts its 
defensive posture (Figure lb). Since the spinal cord 
lies posterior to the vertebral column, extreme flexion 
would place the neural elements (spinal cord and cauda 
equina) under considerable tension, amounting to an 
increase of 15% in length or 6 cm in a large adult. The 
cauda equina in this animal is collectively as thick as 
the spinal cord, but consists of multiple nerve bundles 



Echidna - ambulatory posture 



Spinal Cord 



TV7 



Cauda equina 
and spinal nerves 





Coccygeal 



T 



Mi 



\ % 



t> 



.;vh 



100 Jim vc ^ 



Echidna - defensive posture 

TV7 



Spinal Cord 



Cauda equina 
and spinal nerves 




d 
DC 



NP 







■j< 



100|Lim 



\ X MZ 
SG 




Figure 1. The spinal cord of the Australian echidna is very short, ending opposite the seventh thoracic 
vertebra when the animal is in the ambulatory posture (a). This may be an adaptation which allows 
pronounced flexing of the vertebral column in the defensive posture (b), since the spinal nerves of the 
cauda equina would be more tolerant of the stretching associated with flexing of the vertebral column 
than the much thicker and more vascular spinal cord. Figures c and d show features of the spinal cord 
reported in Ashwell and Zhang (1997). Please see that paper for ethical clearance details and tissue 
preparation methods. Figure lc shows the large neurons of the median nuclear group (arrowhead) at the 
lower lumbar level of the spinal cord (L4). The small inset indicates the position of the larger image. CC 
- central canal of spinal cord: DC - dorsal column; VC - ventral column; VH - ventral horn. Figure Id 
shows unmyelinated afferent fibres labelled with a peroxidase conjugated B4 isolectin from Griffonia 
simplicifolia. The small inset indicates the position of the larger image. These afferents enter via Lissauer's 
zone (LZ) and some descend to deep layers of the dorsal horn (arrowhead) terminating in the nucleus 
proprius (NP), unlike unmyelinated afferents to therian spinal cord, which are confined to the superficial 
layers; e.g. marginal zone - MZ; substantia gelatinosa - SG). 



288 



Proc. Linn. Soc. N.S.W., 125, 2004 



M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 



which are free to move independently of each other, 
unlike the spinal cord where individual axons are 
tightly bound together and are surrounded by delicate 
capillaries. Therefore the stretching of neural elements 
associated with extreme vertebral flexure, which is not 
only large of itself but also affects different segmental 
nerve roots to a greater or lesser extent, is perhaps more 
easily accommodated by shifting more of the nerve 
pathway length into the cauda equina. 

At a histological level, the spinal cord was 
found to have similar cytoarchitectural features 
characterising the laminar organization to that seen in 
the spinal cords of eufherian mammals (Ashwell and 
Zhang 1977). Spinal cord nuclei found in eutherians 
were also identified in the monotreme, except for the 
central cervical nucleus. In addition, a distinct group 
of large neurons, named the median nuclear group, 
was identified in the ventral part of Rexed's lamina X 
and extending into the ventral funiculus at the lower 
lumbar level (Figure lc). Fibre calibre in the dorsal 
and ventral roots of the echidna was similar to that 
reported in eutheria, suggesting similar proportions of 
afferent fibre classes and a and P motorneurons. 

The distribution of unmyelinated primary 
afferent fibres within the dorsal horn of the echidna 
spinal cord have been examined using lectin labelling 
with Griffonia simplicifolia isolectin B4. When 
conjugated with horseradish peroxidase, GSB4 is 
known to label unmyelinated primary afferents 
terminating in both the dorsal horn and cranial nerve 
sensory nuclei (Streit et al. 1985; Plenderleith et al. 
1989; Ashwell and Zhang 1997). It was seen that the 
pattern of labelling with this lectin within the spinal 
cord differed significantly from that seen in eutheria 
in several respects. Firstly, while labelling was seen 
within layers I and II of the echidna dorsal horn (similar 
to eutheria, Streit et al. 1985; Plenderleith et al. 1989), 
labelled fibre bundles were also seen coursing around 
the lateral margin of the dorsal horn as well as through 
layers I and II to terminate in deeper layers of the 
echidna dorsal horn (Figure Id). In eutheria, lectin 
labelled primary afferents terminate only in the 
superficial layers of the dorsal horn (Streit et al. 1985; 
Plenderleith et al. 1989). This deeper labelling in the 
echidna was found to consist of discrete patches in the 
central and lateral parts of layers III and V 
(corresponding to the nucleus proprius). Furthermore, 
in upper cervical segments of the echidna spinal cord, 
labelled axons were identified coursing around the 
margins of the dorsal columns to terminate in the 
internal basilar nucleus (Ashwell and Zhang 1997). 
These two aforementioned features reflect unusual 
primary afferent termination in the echidna, but the 
elucidation of the functional significance of these 
would require electrophysiological studies. Generally 



however, spinal cord cytoarchitectural organization 
seems to be highly conserved across mammals. 



CORTICOSPINAL TRACT 

The echidna corticospinal tract (Figure 2a) 
differs from other mammals (Figure 2b, c, d) in both 
its position within the brainstem and in the level at 
which it decussates (Goldby 1939). The tract runs 
through the cerebral peduncle, decussates in the pons, 
and continues in the lateral medulla, dorsal to the spinal 
tract of the trigeminal nerve. At the spinomedullary 
junction it enters the most posterior part of the lateral 
column of the spinal cord and has been traced as far 
caudally as the 24fh spinal segment, which corresponds 
to lower lumbar to upper sacral levels. No evidence 
has been found for the presence of a pyramidal tract 
close to the ventral midline of the medulla, nor for a 
decussation in the usual position at the caudal end of 
the medulla, as seen in most eutheria. In no other 
mammal is the pyramidal decussation as high as in the 
echidna, nor does the tract, after decussation, lie in 
such an extreme lateral position as in this monotreme. 
It is of interest to note, however, that a high decussation 
of the pyramidal tract is particularly characteristic of 
a small number of highly specialised mammals, which 
probably developed these corticospinal specialisations 
at a very early period in mammalian evolution (Goldby 
1939). For example, some bats and edentates have a 
decussation just caudal to the pons and there is a 
tendency in some of these mammals for fibres from 
this high decussation to take up a lateral position in 
the medulla, e.g. in an armadillo, Lysiurus unicinictus, 
and the pangolin, Manis tricuspis (Goldby 1939). Since 
both of these eutherians are capable of pronounced 
vertebral flexure, as is the echidna, one is tempted to 
speculate that a high pyramidal tract decussation may 
be advantageous for mammals which use this type of 
defensive posture, although the precise nature of the 
advantage which this may confer is not clear at present. 

In polyprotodontid metatheria e.g. the 
American opossum Didelphis virginiana, the 
corticobulbar and corticospinal tracts have been shown 
to be small and probably extend no further than the 
upper cervical segments of the spinal cord (Turner 
1924, see also review by Heffner and Masterton 1983) 
and yet as noted above the pyramidal tract in the 
echidna is much more extensive. Among eutherians, 
both hedgehogs and tree shrews (Figure 2c) show 
termination of the corticospinal tract at higher 
segmental levels (upper cervical for the hedgehog and 
midthoracic for the tree shrew, for review see Heffner 
and Masterton 1983) than that seen in the echidna. 
These observations have made the extensive and 



Proc. Linn. Soc. N.S.W., 125, 2004 



289 



ECHIDNA CENTRAL NERVOUS SYSTEM 



a) Echidna 




b) Marsupial (e.g. Phascolartus 
or Pseudochirus) 



s 
p 

i T1 
n 
a 
I 

c L1 

o S1 

r 

d 



\J 



Medulla 




c) Edentates and Chiroptera 



d) Human 





unusual corticospinal pathway of the 
surviving monotremes of particular 
interest. Extension of the corticospinal 
tract down the greater length of the spinal 
cord is usually regarded as a feature of 
advanced neurological organization, as 
seen in primates (Figure 2d) and 
carnivores, because it allows direct 
control of the cerebral cortex over motor 
units within many levels of the spinal 
cord. 



BRAINSTEM AND 
HYPOTHALAMUS 

The gracile and cuneate nuclei 
are extraordinarily large in the echidna 
(Figure 3a), reflecting the well- 
developed somatosensory pathways for 
the limbs of the echidna. Furthermore, 
estimates of the proportion of white 
matter in the dorsal columns to total 
white matter in the cord gave an average 
result of 25%, suggesting well-developed 
trunk and appendicular somatosensory 
pathways comparable in development to 
carnivores and primates (Ashwell and 
Zhang 1997). In absolute terms the dorsal 
column pathway is as large as that in the 
domestic cat and Macaca fuscata - 
therians of similar body weight. This 
degree of development of the dorsal 
columns ranks the echidna among the 
most neurologically specialized primates 
with well-developed discriminative 
tactile sense. Perhaps the high level of 
somatosensory development can be 
attributed to dense innervation of the 
echidna's forelimb (Mahns et al. 2003) 



Figure 2 Diagrammatic summary of the course, size and extent of the corticospinal tract (bold) in 
representative mammals. Note that the corticospinal tract in the echidna is large, has a high decussation 
and extends to caudal levels of the spinal cord. Contrast this with the small size of the corticospinal tract 
in marsupials (b) and bats and edentates (c) and restriction of the tract to upper segmental levels of the 
spinal cord in those mammals. In size and extent, the echidna corticospinal tract is more like that seen in 
primates (d) and carnivores: mammals in whom a long and large corticospinal tract is believed to confer 
neurological advantages in the form of direct cortical control of motor units in the spinal cord. The 
corticospinal tract of the echidna also has a relatively high level of decussation (crossing over) compared 
to therian mammals, although some mammals (e.g. edentates and chiroptera - c) with the ability to flex 
their vertebral column also have a high level of decussation. No undecussated ventral corticospinal tract, 
as seen in primates (d) has been reported in the echidna. Data for the echidna is derived from Goldby 
(1939), while data for other mammals comes from Kappers, Huber and Crosby (1960). 



290 



Proc. Linn. Soc. N.S.W., 125, 2004 



M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 




Figure 3. Coronal cryostat section (40 um thickness) through the brainstem of an echidna stained for 
cytochrome oxidase by the Wong Riley technique (Wong-Riley 1979)(a, b) and Nissl substance (c, d). 
Please see Hassiotis and Ashwell (2003) for details of experimental ethics and animal acquisition. Strong 
cytochrome oxidase reactivity demonstrates the presence of high densities of mitochondria in axon 
terminals of major sensory pathways for limb and trunk somatosensory pathways (e.g. cuneate nucleus) 
and cranial somatosensory pathway (e.g. nucleus of the trigeminal spinal tract). The mesencephalic nucleus 
of the trigeminal nerve occupies a midline position dorsal to the cerebral aqueduct. The inset in c indicates 
the position of d. 3 - oculomotor nucleus; 4v - fourth ventricle; 12 - hypoglossal nucleus; Aq - cerebral 
aqueduct; Cu - cuneate nucleus; IO - inferior olivary nuclear complex; mcp- middle cerebral peduncle; 
Pn - pontine nuclei; SC - superior colliculus; t5 - trigeminal spinal tract; Vc -caudal part of the nucleus 
of the trigeminal spinal tract ; Ve - vestibular nuclei; Vmes - mesencephalic nucleus of the trigeminal 
nerve; Vo -oralis part of the nucleus of the trigeminal spinal tract. 



or spines, although this has never been studied 
histologically. Alternatively, this specialization may 
have arisen because the echidna spends time in 
subterranean channels, where visual and auditory input 
are of little benefit, and the sense of smell and touch 
are of the most value. At present there are no 
morphological studies of echidna postcranial tactile 



receptors available to shed light on this. 

The trigeminal nerve is also greatly enlarged 
in the echidna as are the nuclei of the trigeminal spinal 
tract (Figure 3a, b). This is consistent with the 
impression from behavioural and electroreception 
studies that the echidna's snout is extremely sensitive 
(see above). The trigeminal system in the echidna 



Proc. Linn. Soc. N.S.W., 125, 2004 



291 



ECHIDNA CENTRAL NERVOUS SYSTEM 



displays a high degree of specialisation similar in kind 
to that seen in Ornithornychus, but not to such a large 
extent. In other words, it does not appear to be as 
sensitive an electroreceptive tool as the platypus bill 
(Proske et al. 1998). Another note-worthy feature is 
that the motor nucleus of the trigeminal nerve in the 
echidna brainstem is much larger than would be 
expected in an animal whose jaw musculature is so 
poorly developed (Abbie 1934). 

The mesencephalic nucleus of the fifth nerve 
in echidna is very like that seen in reptiles in that it 
adopts an almost exclusively mid-line distribution 
(Abbie 1934, Figure 3c, d). Metatheria exhibit a 
condition intermediate between that of the echidna and 
eutheria with more extensive development of the lateral 
mesencephalic V extensions. The mesencephalic 
nucleus and root of the fifth nerve are generally 
considered as being concerned with proprioceptive 
sensibility of jaw musculature. Since the echidna has 
very poor jaw musculature, such a pronounced 
development of mesencephalic V is inexplicable. 

The echidna auditory and vestibular apparatus 
are also notable. In Ornithorhynchus , the entire 
labyrinth has been described as being typically avian 
(Gray 1908). In the echidna, the inner ear shows 
dissimilarities to therians, in that the echidna cochlea 
is banana shaped and has only half a turn, hence is 
partially coiled, whereas in humans the cochlea has 
two and a half turns, and is fully coiled (Gray 1908). 
The cochlea also shows maximal response to sound of 
about 5kHz, substantially lower than in eutheria 
(Augee and Gooden 1993). It has been proposed that 
when the cochlea evolved from the primitive labyrinth, 
it employed the existing vestibular connections within 
the brain, and that when the cochlear apparatus attained 
a mammalian level of structural specialization, a 
trapezoid body appeared in the brainstem. The 
trapezoid body in the echidna is so rudimentary that it 
reveals its primitive vestibular and primarily trigeminal 
origin, because it consists almost entirely of vestibular 
parts and external arcuate fibres which include a large 
trigeminal element (Winkler 1921; Abbie 1934). In 
therians, the pronounced increase in auditory fibres 
almost completely obscures the original trigeminal and 
vestibular connection. Winkler (1921) has argued that 
the poor cochlear development in the echidna renders 
the vestibular fibres relatively conspicuous. While 
central auditory pathways have never been closely 
examined in the echidna, these observations suggest 
that those pathways are either organized differently or 
not as extensive as in theria. 

The hypothalamus in the echidna has been 
reported to have few striking features (Abbie 1934). 
The mammalian hypothalamus is very old 



phylogenetically (Simerly 1995) and very conservative 
in structure throughout the vertebrate series. One 
peculiarity, which links the echidna hypothalamus with 
that of reptiles, and is in sharp contrast to the majority 
of mammals, is the extremely poor development or 
possible absence of the echidna mammillothalamic 
tract (Abbie 1934). Regidor and Divac (1987) for 
example found no evidence of the mammillothalamic 
tract in the echidna on examination of myelin-stained 
coronal sections. This pathway is a key link in the 
Papez circuit underlying memory and emotions in 
eutheria. Its poor development in the echidna may 
indicate that this mammal has an alternative circuit 
for these functions. 



THE ABSENCE OF A CLAUSTRUM IN THE 
FOREBRAIN 

The absence of a claustrum in the echidna 
was initially noted by Abbie (1940) and by Divac and 
co-workers (Divac et al. 1987a) and was further 
discussed more recently (Butler et al. 2002). Similarly, 
no claustrum has been identified in the platypus brain 
(Butler et al. 2002). This structure has been identified 
in all therian mammals so far examined (Johnson et 
al. 1994) and is believed to have a structural and 
chemical affinity with the neocortex, although its 
precise functional significance is uncertain. It engages 
in reciprocal connections with neocortex and receives 
projections from the non-specific intralaminar nuclei 
of the thalamus (see Butler et al. 2002 for review). 
The question remains open as to whether the claustrum 
was present in ancestral mammals and disappeared in 
the monotremes, or whether its evolution represents 
an exclusively therian brain development. 



THE DISTRIBUTION OF CHEMICALLY 
IDENTIFIED NEURONS 

Manger and co-workers have recently 
examined the distribution of cholinergic, 
catecholaminergic and serotonergic neurons in the 
brains of the platypus and echidna (Manger et al. 
2002a, b, c). Those authors showed that while there 
are many similarities between monotremes and therians 
in the distribution of these neurons, there were also 
some evolutionarily and potentially functionally 
significant differences. For example, cholinergic cells 
are present in the monotreme brain, but important cells 
groups identified in theria do not appear to be present 
in the platypus or echidna. These include cholinergic 
cells in the cerebral cortex, nuclei of the vertical and 



292 



Proc. Linn. Soc. N.S.W., 125, 2004 



M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 




Figure 4. Coronal cryostat sections (40 urn thickness) through the caudal thalamus (a) and rostral thalamus 
(b) stained for cytochrome oxidase and Nissl substance, respectively. Please see Hassiotis and Ashwell 
(2003) for details of experimental ethics and animal acquisition. Note the large ventral posterior thalamic 
nucleus with lateral (VPL) and medial (VPM) compartments. In theria these two regions serve processing 
of somatosensory (touch) information from the body and head, respectively. Contrast the size of these 
two nuclei with the lateral geniculate nucleus (LG) processing visual information. The reticular thalamic 
nucleus, which is found in all therian mammals external to the ventral tier nuclei (i.e. embedded in the 
external medullary lamina to the left of 4a), appears to be absent from the echidna thalamus. The rostral 
thalamus (b) contains a large nucleus (anteromediodorsal -AMD) with no clear division into subnuclei. 
This may correspond to the mediodorsal nucleus of therians. 3v - third ventricle; eml - external medullary 
lamina; ml - medial lemniscus; ZI - zona incerta. 



horizontal limbs of the diagonal band of Broca, the 
magnocellular preoptic nucleus, the substantia 
innominata, nucleus of the ansa lenticularis, 
hypothalamic nuclei and the parabigeminal nucleus 
(Manger et al. 2002a). They proposed that the absence 
of cholinergic neurons from the hypothalamus might 
be related to the unusual features of monotreme sleep 
(Siegel et al. 1996, 1998). 

The catecholaminergic system of the 
monotreme brain appears to be very similar to that 
found in theria, but there were some minor differences 
in the form of the absence of A4, A3 and C3 groups 
from the locus coeruleus and caudal rhombencephalon. 
It should be noted however, that these are only small 
differences and this great similarity demonstrates the 
high degree of evolutionary conservatism in these 
neurons across amniote species (Manger et al. 2002b). 



Serotonergic neurons in monotremes appear 
to fall into three groups: hypothalamic, rostral nuclear 
and caudal nuclear clusters. The rostral and caudal 
nuclear groups are found consistently across all 
mammals while the hypothalamic cluster, although not 
reported in other mammals, is found in most other 
species of vertebrates (Manger et al. 2002c). 



THE THALAMUS AND THALAMOCORTICAL 
PROJECTIONS 

Campbell and Hayhow (1971) identified 
several thalamic nuclei in echidna, which exhibited 
cyto- and myeloarchitectonic features resembling those 
found in other mammals (Figure 4). However, echidna 
thalamic nuclei are not as easily distinguished as those 



Proc. Linn. Soc. N.S.W., 125, 2004 



293 



ECHIDNA CENTRAL NERVOUS SYSTEM 



in opossums (Bodian 1939, 1942; Oswaldo-Cruz and 
Rocha-Miranda 1968; Benevento and Ebner 1971) or 
other commonly used laboratory mammals (Rose 
1942; Rose and Woolsey 1949). Chemoarchitectural 
characteristics of the thalamus in echidnas and rats have 
been compared in sections stained for myelin, 
acetylcholinesterase (AChE), succinate dehydrogenase 
(SDH) and cytochrome oxidase (CO) by Regidor and 
Divac (1987). Numerous species differences were 
noted, but in general the thalamus is architecturally 
more homogenous in echidnas than in rats, especially 
within the anterior portion (Figure 4b). The large 
structure localized in the anteromediodorsal part of the 
thalamus of the echidna has been found to contain small 
amounts of acetylcholinesterase and oxidative 
enzymes; in this respect resembling the mediodorsal 
nucleus of rats. Regidor and Divac (1987) concluded 
that this brain structure of echidnas corresponds to the 
mediodorsal nucleus in placental species. 

Welker and Lende (1980) defined and 
described the thalamic nuclei that contribute major 
projections to the isocortex in echidna. Their purpose 
was to determine whether the echidna thalamus 
exhibited mammalian thalamocortical relations more 
similar to those found in metatheria, or to those in 
eutherian mammals. Welker and Lende also attempted 
to identify whether an enlarged thalamic nucleus was 
sending afferents to the enlarged frontal cortex. They 
performed a series of partial ablations of the somatic 
sensory, auditory, visual and motor areas, as well as 
in several different portions of the greatly enlarged 
frontal neocortex (see below) and demonstrated that 
the thalamocortical connections in the echidna are 
similar in most respects to those demonstrated in 
eutherian mammals. One unusual feature observed by 
Welker and Lende was a large nuclear mass in the 
dorso-fronto-medial thalamus (presumptive 
anteromediodorsal nucleus discussed above), which 
projects to the enlarged frontal cortex (Divac et al. 
1987a, b). It has been hypothesised that this nuclear 
region is homologous to the eutherian mediodorsal 
nucleus. Their data also revealed that projections to 
separate motor and somatic sensory cortical areas from 
the thalamus were spatially distinct (Welker and Lende 
1980). 



CORTICAL STRUCTURE AND 
ORGANIZATION 

Until the late 1 800's it was generally believed 
that all mammals possessed a corpus callosum (Turner 
1 890). a major fibre bundle connecting the neocortex 
of the two hemispheres of the brain. Elliott Smith 



(1902, 1903) dispelled this notion in his early studies 
of comparative cortical organization. He showed that 
in monotremes and metatheria, the anterior 
commissure is the major cerebral commissure, being 
the sole connection between all parts of the neo- and 
paleocortex, with only a small archicortical 
commissural connection being present dorsally (the 
hippocampal commissure). 

Several striking aspects of gross cortical 
anatomy have been noted in Tachyglossus aculeatus. 
The most obvious of these is the high degree of 
gyrification (36% of isocortex buried in fissures), 
comparable to that in many eutherian mammals (e.g. 
cat 40%, squirrel monkey 39%). The second is the large 
proportion of the brain volume occupied by the 
cerebral cortex (43%), similar to values in eutheria 
(prosimians - 54%, Pirlot and Nelson 1978). Among 
the brains of eutheria, a highly gyrified cerebral cortex 
is usually considered as an attempt to maximise the 
number of cortical columns available for the processing 
of information. Therefore a highly gyrified cortex is 
considered the hallmark of more neurologically 
advanced mammals such as carnivores, primates and 
cetaceans. This raises the question as to why an animal 
like the echidna, which leads a solitary existence and 
has no known complex social life, has such a highly 
gyrified cortex. One principal difference between the 
brains of the two living Australian monotremes is that 
the platypus' cortex is quite smooth (lissencephalic), 
whereas the echidna cortex is complexly folded. 

Another most remarkable aspect of echidna 
neurobiology concerns cortical topography. Ziehen 
(1897, 1908), Brodmann (1909) and Schuster (1910), 
all published early observations on the cortex in the 
Monotremata. Brodmann examined the cortex in 
echidna and established that there is a typical six- 
layered distribution. The echidna has been noted to 
have a thinner cortex, perhaps due to its denser packing 
of neurons compared to the platypus (Abbie 1940). In 
both the echidna and the platypus, Ziehen ( 1 897, 1 908) 
showed that there was a change in the type of cortex 
between the anterior (olfactory) and posterior 
(sensorimotor) portions of the hemisphere, and 
Schuster (1910) confirmed his observations. 
Nevertheless, these early authors concluded that the 
plan upon which the monotreme brain is constructed 
conforms in every respect to the basic pattern 
prevailing among the vast majority of other mammals 
(Abbie 1940). 

To date, the most detailed anatomical study 
of the echidna cortex was performed in the 1940's by 
Abbie. Since the 1940's, no further in-depth anatomical 
studies have been done on the anatomy of the echidna 
cortex as a whole, although specific systems have been 



294 



Proc. Linn. Soc. N.S.W., 125, 2004 



M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 



Welker and Lende (1980) 




Figure 5. Electrophysiology and cytoarchitecture of the echidna cerebral cortex. The earliest study 
illustrated is by Lende (1964)(results shown in a summary diagram redrawn from Welker and Lende 
1980). Greek letters denote major consistent sulci as delineated by Smith (1902). The Welker and Lende 
map shows only the externally visible gyral surface and indicates the position of motor (M), somatosensory 
(SI), visual (V) and auditory (A) cortices. The Ulinski map shows cytoarchitectural fields identified by 
that author (Ulinski 1984). Two coronal sections (i and ii) are illustrated with the positions of rostral (r) 
and caudal (c) fields of the somatosensory cortex (SMI) marked. The small lateral view shows the 
rostrocaudal positions from which these sections were taken. The figure below the two sections shows a 
flattened representation of cortex. Note that Ulinski's "r" field lies rostral (and superior) to the deepest 
part of the a sulcus. The lower two illustrations summarize the findings of Krubitzer et al (1995). The 
smaller diagram shows a representation of the entire flattened cortical surface with the sulcal walls opened. 
Solid lines in the Krubitzer map indicate the deepest point of the sulci, while dotted lines indicate the 
sulcal rims at the external cortical surface. The larger illustration shows a drawing of the completely 
flattened cortical surface with the boundaries of functional areas indicated relative to P and a sulci (thick 
grey lines). Note that the rostral somatosensory field in Krubitzer's map lies caudal to the a sulcus (cf 
Ulinski map). Ent - entorhinal cortex; Hi - hippocampus; M - manipulation cortex; Pir - piriform cortex; 
PV - parietal ventral somatosensory cortex; R -rostral somatosensory cortex; SI - primary somatosensory 
cortex. 



Proc. Linn. Soc. N.S.W., 125, 2004 



295 



ECHIDNA CENTRAL NERVOUS SYSTEM 



studied. Abbie described the monotreme neocortex as 
comprising two broad components; one related to the 
hippocampus, labelled by him as parahippocampal 
regions and located in the anterior and medial parts; 
and the other related to the piriform cortex, labelled as 
the parapiriform regions, located posteriorly and 
laterally. He also defined sulcal boundaries to these 
regions. When labelling the cortex, Abbie adopted the 
system of Elliot Smith (1902), using Greek letters to 
name the major and deepest sulci (Figure 5). There 
are two pronounced sulci in the monotreme cortex, 
denoted as a and p\ These divide the frontal cortex 
from the posterior motor and sensory cortices. 

More recent functional studies of the 
isocortex of Tachyglossus aculeatus have indicated that 
the primary motor, somatosensory, auditory and visual 
areas are located in the caudal half of the isocortex 
(Lende 1964)(Figure 5). Aside from the posterior 
location of these areas, the following relationships are 
unlike those described in any other therian mammals: 
the somatic sensory area is confined to the ventral 
portion of the lateral surface; the visual area is located 
dorsal to the somatic sensory area and borders the 
representation for the tail; the auditory area is located 
posterior to the visual and somatic sensory areas and 
borders the latter at the representation of the back. 
These relationships might be described as rotational 
dislocation of the areal relations relative to that found 
in eutherians in that the somatic sensory area has been 
displaced downward and backward, the auditory area 
upward, and the visual area upward and forward 
(Lende 1964). 

The somatosensory representation in the 
echidna is in some respects similar to that of other 
mammals. The area for the tail is found uppermost 
and the areas for hind limb, trunk, forelimb, and head 
are located laterally and ventrally, in that sequence. 
This is the same as the basic mammalian pattern of 
somatosensory area 1 (SI) as established by Woolsey 
(1952). A relatively large portion of somatosensory 
cortex in the echidna was found by Lende to be devoted 
to the head, and particularly the snout and tongue, as 
might be expected from the ant-eating habits of the 
echidna (Lende 1964). 

Physiological studies have indicated that 
more than 50% of the rostral cortex of the echidna has 
no attributable primary motor or sensory function and 
has been considered as an expanded prefrontal cortex 
(Welker and Lende 1980). If this interpretation is 
correct, then the proportion of isocortex in 
Tachyglossus aculeatus occupied by the prefrontal area 
exceeds that in humans (29%) Divac et al. (1987a, 
b)(see section on thalamus and thalamocortical 
projection in previous pages). 

Ulinski's study (1984) examined the 



cytoarchitecture and thalamic afferents of the 
somatosensory area (SMI) in the echidna. His findings 
indicated that SMI contains two cytoarchitectonic 
fields. A caudal field with a well-developed layer IV 
present extends across the post a gyrus and onto the 
floor of sulcus a The rostral field was reported to 
extend from the floor of sulcus a onto its rostral bank. 
It also was reported to have a well-developed layer IV 
but with a large number of pyramidal neurons in layer 
V. The remainder of the pre a gyrus was reported to 
contain a single cytoarchitectonic field with a thin layer 
IV and layer V heavily populated with larger pyramidal 
cells. This field corresponded to the physiologically 
defined motor area Ml. Thalamic afferents to 
somatosensory area were examined by placing pressure 
injections of horseradish peroxidase into the two 
architectonic fields. The results indicated that the 
somatosensory area in Tachyglossus aculeatus contains 
two cytoarchitectonic fields that resemble areas 3a and 
3b in some placental mammals, leading Ulinski to the 
conclusion that the collection of cytoarchitectonic 
fields corresponding to areas 4, 3a, and 3b is a basic 
mammalian character which has been modified in 
metatherian and many eutherian mammals. 

In more recent times, Krubitzer et al. (1995) 
undertook a detailed study of monotreme cortical 
organization as part of a comparative approach to 
determining those features of the isocortex which 
characterise all the major lines of mammalian 
evolution, More specifically, their investigation was 
designed to determine the internal organization and 
number of somatosensory fields in the monotreme 
isocortex. 

The isocortices of both monotremes were 
found to contain four representations of the body 
surface. A large area that contained neurons 
predominantly responsive to cutaneous stimulation of 
the contralateral body surface was identified as the 
primary somatosensory area (SI). This was found 
caudal and ventral to the a sulcus. Another 
somatosensory field (R) was identified rostral to SI. 
The topographic organization of R was similar to that 
found in SI, but neurons in R were responsive most 
often to light pressure and taps to peripheral body parts. 
Neurons in cortex located rostral to R were responsive 
to manipulation of joints and hard taps to the body. 
This field was termed the manipulation field (M) and 
occupies the position of the motor cortex identified 
by Lende. Note that Krubitzer' s M field occupies an 
area which Ulinski denoted as the rostral 
somatosensory field and Krubutzer's R field occupies 
at least part of Ulinski's caudal somatosensory field 
(Ulinski 1984)(Figure 5). Consequently the two studies 
are not easily reconciled. A parieto-ventral 
somatosensory field (PV) was also identified by 



296 



Proc. Linn. Soc. N.S.W., 125, 2004 



M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 




I 



^V' 



i 



M 



SI 



*>-* 



IV 






dorsal ax£ 

si>" .R. 



"500 urn 




100 urn 



Figure 6. NissI stained cryostat sections in the coronal plane through the cerebral cortex and olfactory 
bulb of the echidna. Please see Hassiotis and Ashwell (2003) for details of experimental ethics and animal 
acquisition. The inset drawings show: i) a lateral view of the echidna cerebral hemisphere indicating the 
planes of section shown in a, b, c and d, e, respectively; and ii) a line drawing of a coronal section showing 
the positions from which a, b, and c were taken. Figure 6a shows a lower power view of motor cortex (M) 
and the rostral field of somatosensory cortex (R). Figures 6b and c show motor cortex and SI somatosensory 
cortex, respectively with layers indicated by Roman numerals. WM - subcortical white matter. Figures 
6d and e show low and high power views of the olfactory bulb. Rectangle in d indicates the position of e. 
Note the lack of a clear tightly grouped monolayer of mitral cells (Mi) as is seen in therian mammals 
(Switzer and Johnson 1977). E - ependyma of lateral ventricle; Gl - glomerular layer; IGr - internal 
granular layer; IPL - internal plexiform layer; ON - olfactory nerve fibre layer; Pir - piriform cortex. 



Proc. Linn. Soc. N.S.W., 125, 2004 



297 



ECHIDNA CENTRAL NERVOUS SYSTEM 



Krubitzer and was thought to be homologous to its 
therian counterparts (Krubitzer et al. 1995). The 
evidence for the existence of four separate 
somatosensory representations in somatosensory 
cortex was taken to indicate that cortical organization 
is more complex in the echidna than had been 
previously thought. Furthermore, although the two 
monotreme families have been quite separate for at 
least 55 million years (Richardson 1987), the similarity 
of cortical field organization in both monotremes 
studied suggested either that the original differentiation 
of sensory fields occurred very early in mammalian 
evolution, or that the potential for division of 
somatosensory cortex into numerous fields was highly 
constrained in evolution, so that both species arrived 
at the same result independently. 

Figure 6 shows the cytoarchitecture of the 
echidna motor and somatosensory cortices. 
Nomenclature for cortical areas is adopted from 
Krubitzer et al (1995). As in eutherian motor cortex 
(Figure 6b), the echidna M cortex is characterised by 
large pyramidal neurons in layer V. The SI part of 
somatosensory cortex (Figure 6c) is characterised by 
a layer IV rich in densely packed small neurons. 

Several other aspects of cortical function in 
this species are also of note; particularly the apparent 
absence of an SII somatosensory representation and 
the relatively lateral position of the motor 
representation compared to that in eutheria (Krubitzer 
et al. 1995). Until recently, functional studies had failed 
to identify any parietal association cortex, but 
Krubitzer et al. (1995) was able to identify a 
topographically discrete, multimodal area between the 
primary sensory cortical areas, which may represent 
such an area. 



CONCLUDING REMARKS 

There are a number of unusual features of 
the anatomy of the brain and spinal cord of the echidna. 
Some of these are probably indicative of the early 
branching of the monotreme lineage from therian 
mammals (e.g. absence of the claustrum), some have 
potential functional significance for the unusual 
physiology of this mammal (e.g. the absence of 
hypothalamic cholinergic neurons), while some appear 
to be peculiar adaptations to the niche occupied by 
this mammal (e.g. trigeminal nuclei development and 
short spinal cord). 



ACKNOWLEDGEMENTS 

This work was supported in part by a grant from the 
Australian Research Council. We are grateful for the 
assistance of Drs Luan ling Zhang and Hong qin Wang 
in the preparation of the Nissl, cytochrome oxidase 
and lectin stained sections through the echidna brain 
and spinal cord. 



REFERENCES 

Abbie, A.A. (1934). The brainstem and cerebellum of 

Echidna aculeata. Philosophical Transactions 
of the Royal Society of London Series B 
Biological Sciences 224, 1-74. 

Abbie, A.A. (1940). Cortical lamination in the 
monotremata, Journal of Comparative 
Neurology 72, 429-467. 

Andres, K.H, von During M., Iggo, A. and Proske, U. 
(1991). The anatomy and fine structure of the 
echidna Tachyglossus aculeatus snout with 
respect to its different trigeminal sensory 
receptors including the electroreceptors, 
Anatomy and Embryology 184, 371-393. 

Ashwell, K.W.S and Zhang, L.L. (1997). Cyto- and 

myeloarchitectonic organization of the spinal 
cord of an echidna (Tachyglossus aculeatus), 
Brain, Behaviour and Evolution 49, 276-294. 

Augee, M and Gooden, B. (1993). 'Echidnas of Australia 
and New Guinea' . (New South Wales 
University Press, Sydney). 

Benevento, L.A and Ebner, F.F. (1971) The contribution 
of the dorsal lateral geniculate nucleus to the 
total pattern of thalamic terminations in striate 
cortex of the Virginia opossum. Journal of 
Comparative Neurology 143, 243-260. 

Bodian, D. (1939) Studies on the diencephalon of the 
Virginia opossum; the nuclear pattern in the 
adult. Journal of Comparative Neurology 71, 
259-323. 

Bodian, D. (1942) Studies on the diencephalon of the 
Virginia opossum; the thalamocortical 
projection. Journal of Comparative Neurology 
77, 525-575 . 

Butler, A.B., Molnar, Z., and Manger P.R. (2002) 

Apparent absence of claustrum in monotremes: 
Implications for forebrain evolution in amniotes. 
Brain, Behaviour and Evolution 60, 230-240. 

Brodmann, K. (1909). Vergleichende Lokalisationslehre 
- Lehre der Grosshirnrinde in ihren Prinzipien 
dargestellt auf Grand des Zellen baues' (Barth, 
Leipzig). 

Campbell, C.B. and Hayhow, W.R. (1971). Primary optic 
pathways in the echidna Tachyglossus 
aculeatus: an experimental degeneration study, 
Journal of Comparative Neurology 143, 1 1 9- 
136. 



298 



Proc. Linn. Soc. N.S.W., 125, 2004 



M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 



Divac, I, Hoist, M.C, Nelson, J and McKenzie, J.S. 

(1987a). Afferents of the frontal cortex in the 
echidna (Tachyglossus aculeatus): indication of 
an outstandingly large prefrontal area, Brain, 
Behaviour and Evolution 30, 303-320. 

Divac, I, Pettigrew, J.D, Hoist, M.C and McKenzie, J.S. 
(1987b). Efferent connections of the prefrontal 
cortex of echidna (Tachyglossus aculeatus), 
Brain, Behaviour and Evolution 30, 321-327. 

Goldby, F. (1939). An experimental investigation of the 
motor cortex and pyramidal tract of Echidna 
aculeata. Journal of Anatomy 73, 509-524. 

Gray, A. A. (1908). An investigation on the anatomical 

structure and relationships of the labyrinth in the 
reptile, the bird, and the mammal. Proceedings 
of the Royal Society of London Series B 
Biological Sciences 80, 507-528. 

Gregory, J.E, Iggo, A, Mclntyre, A.K and Proske, U. 

(1987). Electroreceptors in the platypus. Nature 
326, 386-387. 

Gregory, J.E, Iggo, A, Mclntyre, A.K and Proske, U. 
(1988). Receptors in the bill of the platypus. 
Journal of Physiology 400, 349-366. 

Gregory, J.E, Iggo, A, Mclntyre, A.K and Proske, U. 
(1989). Responses of electroreceptors in the 
snout of the echidna Journal of Physiology 414, 
521-538. 

Hassiotis, M. and Ashwell K.W.S. (2003). Neuronal 
classes in the isocortex of an echidna 
(Tachyglossus aculeatus). Brain, Behaviour and 
Evolution 61, 6-27. 

Heffner, R.S and Masterton, R.B. (1983). The role of the 
corticospinal tract in the evolution of human 
digital dexterity. Brain, Behavior and Evolution 
23, 165-183. 

Iggo, A, Mclntyre A.K, Proske U. (1985). Responses of 
mechanoreceptors and thermoreceptors in skin 
of the snout of the echidna Tachyglossus 
aculeatus. Proceedings of the Royal Society of 
London Series B Biological Sciences 223, 261- 
278. 

Johnson, J.I., Kirsch, J.A., Reep, R.L., Switzer, R.C. 3rd. 
(1994). Phylogeny through brain traits: more 
characters for the analysis of mammalian 
evolution. Brain Behaviour and Evolution 43, 
319-47. 

Kappers, C. U. A., Huber, G. C. and Crosby, E. C. (1960). 
'The comparative anatomy of the nervous 
system of vertebrates, including man' . Vol. 1 
(Hafner, New York). 

Krubitzer, L. (1998). What can monotremes tell us about 
brain evolution? Philosophical Transactions of 
the Royal Society of London Series B Biological 
Sciences 353, 1127-1146. 

Krubitzer, L, Manger, P, Pettigrew, J and Calford, M. 

(1995). Organization of somatosensory cortex in 
monotremes: in search of the prototypical plan. 
Journal of Comparative Neurology 351, 261- 
306. 



Lende, R.A. (1964). Representation in the cerebral cortex 
of a primitive mammal: sensorimotor, visual, 
and auditory fields in the echidna (Tachyglossus 
aculeatus). Journal of Neurophysiology 27, 37- 
48. 

Manns, D.A., Coleman, G.T., Ashwell, K.W., Rowe M.J. 
(2003). Tactile sensory function in the forearm 
of the monotreme Tachyglossus aculeatus. 
Journal of Comparative Neurology 459, 173- 
185. 

Manger, P.R and Hughes, R.L. (1992). Ultrastructure and 
distribution of epidermal sensory receptors in 
the beak of the echidna (Tachyglossus 
aculeatus), Brain, Behaviour and Evolution 40, 
287-296. 

Manger, P.R., Fahringer, H.M., Pettigrew, J.D., and 
Siegel, J.M. (2002a). The distribution and 
morphological characteristics of cholinergic 
cells in the brain of monotremes as revealed by 
ChAT immunohistochemistry. Brain, Behaviour 
and Evolution 60, 275-297. 

Manger, P.R., Fahringer, H.M., Pettigrew, J.D., and 
Siegel, J.M. (2002b). The distribution and 
morphological characteristics of 
catecholaminergic cells in the brain of 
monotremes as revealed by tyrosine hydroxylase 
immunohistochemistry. Brain, Behaviour and 
Evolution 60, 298-314. 

Manger, P.R., Fahringer, H.M., Pettigrew, J.D., and 
Siegel, J.M. (2002c). The distribution and 
morphological characteristics of serotonergic 
cells in the brain of monotremes. Brain, 
Behaviour and Evolution 60, 315-332. 

Oswaldo-Cruz, E and Rocha-Miranda, C.E. (1968). The 
brain of the opossum (Didelphis marsupialis): A 
cytoarchitectonic atlas in stereotaxic 
coordinates'. (Instituto de Biofisica, Univ. Fed. 
do Rio de Janeiro). 

Pirlot, P and Nelson, J. (1978). Volumetric analyses of 

monotreme brains, Australian Zoology 20, 171- 
179. 

Plenderleith, M.B., Cameron, A.A., Key, B., Snow , P.J. 
(1989). The plant lectin soybean agglutinin 
binds to the soma, axon and central terminals of 
a subpopulation of small diameter primary 
sensory neurons in rat and cat. Neuroscience 31, 
683-695. 

Proske, U, Gregory, J.E and Iggo, A. (1998). Sensory 
receptors in monotremes. Philosophical 
Transactions of the Royal Society of London 
Series B Biological Sciences 353, 1 187-1 198. 

Richardson, B.J. (1987). A new view of the relationships 
of Australian and American marsupials. 
Australian Mammalogy 11, 71-73. 

Regidor, J and Divac, I. (1987). Architectonics of the 
thalamus in the echidna (Tachyglossus 
aculeatus): search for the mediodorsal nucleus, 
Brain, Behaviour and Evolution 30, 328-341. 

Rose, J.E. (1942) The ontogenetic development of the 

rabbit's diencephalon. Journal of Comparative 
Neurology 77, 61-129. 



Proc. Linn. Soc. N.S.W., 125, 2004 



299 



ECHIDNA CENTRAL NERVOUS SYSTEM 



Rose, J.E. and Woolsey, C.N. (1949). Organization of the 
mammalian thalamus and its relationship to the 
cerebral cortex. Electroencephalography and 
Clinical Neurophysiology 1, 391-404. 

Scheich, H, Langner, G, Tidemann, C, Coles, R.B, 
Guppy, A. (1986). Electroreception and 
electrolocation in platypus. Nature 319, 401- 
402. 

Schuster, E. (1910). Preliminary note upon the cell 

lamination of the cerebral cortex of echidna, 
with an enumeration of the fibres in the cranial 
nerves. Proceedings of the Royal Society of 
London Series B Biological Sciences 82, 1 1 3- 
• 123. 

Siegel, J.M. Manger, P.R., Nienhuis, R., Fahringer, H.M., 
Pettigrew, J.D. (1996) The echidna 
Tachyglossus aculeatus combines REM and 
non-REM aspects in a single sleep state: 
Implications for the evolution of sleep. J. 
Neuroscience 16, 3500-3506. 

Siegel, I.M. Manger, P.R., Nienhuis, R., Fahringer, H.M., 
Pettigrew, I.D. (1998) Monotremes and the 
evolution of rapid eye movement sleep. 
Philosophical Transactions of the Royal Society 
of London Series B 353, 1147-1157. 

Simerly, R.B. (1995). Anatomical substrates of 

hypothalamic integration. In 'The Rat Nervous 
System'. (Ed. G. Paxinos) pp. 353-376. 
(Academic Press, San Diego). 

Smith, G.E. (1902). 'Descriptive and illustrative catalogue 
of the physiological series', in The Museum of 
the Royal College of Surgeons of England, vol 
II, 2 nd edn, Taylor and Francis, Red Lion Court, 
London, pp. 138-157 

Smith, G.E. (1903) On the morphology of the cerebral 
commissures in the Vertebrata, with special 
reference to an aberrant commissure in the brain 
of certain reptiles, Transactions of the Linnean 
Society (London) 2 nd Series Zoology, 8, 455- 
500. 

Streit, W.I., Schulte, B.A., Balentine, I.D., Spicer, S.S. 

(1985). Histochemical localization of galactose- 
containing glycoconjugates in sensory neurons 
and their processes in the central and peripheral 
nervous system of the rat. Journal of 
Histochemistry and Cytochemistry 33, 1042- 
1052. 

Switzer, R.C. 3 rd and Johnson, J.I. Jr (1977) Absence of 
mitral cells in monolayer in monotremes. 
Variations in vertebrate olfactory bulbs. Acta 
Anat (Basel) 99, 36-42. 

Turner, W. (1890) The convolutions of the brain; a study 
in comparative anatomy, Journal of Anatomy 
25. 105-153. 

Turner, E.L. (1924) The pyramidal tract of the Virginian 
opossum (Didelphys virginiana), Journal of 
Comparative Neurology, 36, 387-397. 

Ulinski. P.S. (1984) Thalamic projections to the 
somatosensory cortex of the echidna 
(Tachyglossus aculeatus). Journal of 
Comparative Neurology 229, 153-170. 



Welker, W and Lende, R.A. (1980) 'Thalamocortical 
Relationships in Echidna (Tachyglossus 
aculeatus)' , in Comparative Neurology of the 
Telencephalon, (ed S.O.E Ebbesson) pp. 449- 
481 (Plenum Press, New York). 

Winkler, C. (1921) External arcuate fibers, Encephale 16, 
273-282. 

Wong-Riley, M. (1979) Changes in the visual system of 
monocularly sutured or enucleated cats 
demonstrable with cytochrome oxidase 
histochemistry. Brain Research 171, 11-28. 

Woolsey, C.N. (1952) 'Patterns of localization in sensory 
and motor areas of the cerebral cortex' , in 
Biology of Mental Health and Disease, New 
York, Hoeber, pp. 193-206. 

Ziehen, T.H. (1897) 'Das centralnervensystem der 

monotremen und marsupialier', Makroskopische 
Anatomie, Semon's Zoologische Forschungs, Bd 
3, Teil 1,S 1-187. 

Ziehen, T.H. (1908) 'Das centralnervensystem der 

monotremen und marsupialier', Mikroskopische 
Anatomie, Semon's Zoologische Forschungs, Bd 
3, Teil 2, S 789-921. 



300 



Proc. Linn. Soc. N.S.W., 125, 2004 



Monotreme Tactile Mechanisms: From Sensory Nerves To 

Cerebral Cortex 

Mark J. Rowe, D.A. Mahns and V. Sahai 
School of Medical Sciences, The University of New South Wales, Sydney 2052, Australia 



Rowe, M.J., Mahns, D.A. and Sahai, V. (2004). Monotreme tactile mechanisms: from sensory nerves to 
cerebral cortex. Proceedings of the Linnean Society of New South Wales 125, 301-317. 

Electrophysiological recordings from single tactile sensory nerve fibres supplying the limb extremities in 
the echidna (Tachyglossus aculeatus) reveal a remarkable resemblance between monotreme peripheral tactile 
mechanisms and those of placental mammals. The similarities apply to a concatenation of attributes, including 
the classification of sensory fibre types and aspects of functional properties and tactile coding capacities. 
The analysis demonstrates that high-acuity tactile signalling from the distal forelimb in the monotreme is 
based upon a triad of major tactile fibre classes as is the case for placental mammals. Furthermore, the 
functional similarity between corresponding classes in monotreme and placental species suggests that 
peripheral tactile sensory mechanisms are highly conserved across evolutionarily-divergent mammalian 
orders. 

Evidence for a unique and striking dependence upon tactile sensory mechanisms in monotremes comes 
from both behavioural observations on the animals and from the exceptional prominence given to the 
representation of tactile inputs in the cerebral cortex of these species. In the platypus, for example, almost 
half of its lissencephalic cortex is allocated to the processing of inputs from the bill. Furthermore, within the 
specialized area of bill representation in the platypus cortex, the receptive fields of individual neurones are 
amongst the smallest ever recorded within the somatosensory areas of cortex (often <lmm in diameter), 
presumably conferring great fidelity and precision on the tasks of tactile localization and discrimination 
involving the bill. However, in both the platypus and the echidna there is a complete and highly ordered 
somatotopic representation of tactile inputs from the contralateral body surface, conforming with the so- 
called primary somatosensory cortex (SI) of other mammals. Controversy applies to the issue of whether 
additional, multiple body representations are present in the monotreme cortex, as neither Lende (1964) nor 
Bohringer and Rowe (1977) found evidence of this, in contrast to Krubitzer et al. (1995a), who have argued 
for four body representations in the cortex of both the echidna and the platypus. 

Manuscript received 24 September 2003, accepted for publication 22 October 2003. 

KEYWORDS: echidna, evolution and sensory nerve function, monotreme, Ornithorynchus anatinus, 
platypus, somatosensory system, Tachyglossus aculeatus, tactile receptors, tactile sensory function. 



EVOLUTIONARY PLACE OF THE 

MONOTREME BRAIN thought to represent the next stage of evolution towards 

the Eutherian, or complete mammals. This hierarchical 

The emergence, 100-200 million years ago, concept of the relations between the three great orders 

of monotremes on a separate evolutionary line from f mam mals was, in part, influenced by the presence 

their placental and marsupial mammalian counterparts i n living monotremes of reptilian or avian-like 

has contributed, in particular in the 19 th century, to the reproductive mechanisms. This possession of 

hypothesis that living monotremes are closer to oviparity, or the capacity to lay eggs, set the 

ancestral mammalian forms than their placental and monotremes aside from mammals of the marsupial and 

marsupial relatives, and therefore that the monotreme eutherian orders. Furthermore, a concatenation of other 

nervous system may provide a window on the status anatomical or functional attributes shared with reptilian 

of the ancestral mammalian brain. These notions were an d av i m representatives, but not with other mammals, 

implicit in the designation of monotremes as the te nded to re-inforce these hierarchical concepts. These 

Prototheria, or first mammals, while the marsupial attributes have been documented by Augee and 

mammals were designated the Metatheria and were Gooden (1993) and include, among others, the 



MONOTREME TACTILE MECHANISMS 



arrangement of the shoulder girdle, the less elaborate 
coiling of the cochlea, and the presence of dwarf 
nephrons in the kidney. Perhaps surprisingly, the 
presence of the single opening from the cloaca for both 
reproductive and excretory activity, which led to their 
monotreme designation, is not entirely diagnostic for 
this mammalian group as some marsupials and other 
vertebrates share this feature (Augee and Gooden 
1993). 

Despite the retention of several plesiomorphic 
attributes in living monotremes there should be no 
expectation that such attributes need be a generalized 
feature of this mammalian order. Thus, in other respects 
of their anatomy or physiology, monotremes need not 
have proved any more conservative or constrained in 
their evolutionary progress than their marsupial or 
placental relatives with whom they have shared 
perhaps 200 million years in which to evolve from the 
ancestral forms of their evolutionary forebears. Indeed, 
we see some hint of this evolutionary metamorphosis 
and divergence within the monotreme ranks 
themselves, even in gross features of central nervous 
system organization. For example, although the 
divergence of the monotreme line into separate 
platypus and echidna strands has occurred -50-80 
million years ago (Griffiths 1978; Dawson 1983; 
Richardson 1987), or even as recently as -20 million 
years ago (Belov and Hellman 2003), one encounters 
striking differences between them in the gross 
morphology of the cerebral cortex. The echidna has a 
quite elaborately folded, or gyrencephalic cerebral 
cortex, whereas, in contrast, the platypus possesses a 
smooth, essentially unfolded, lissencephalic cortex. 
The elaborate pattern of the cortical fissures in the 
echidna was analyzed and classified by Elliot Smith 
(1899, 1902) according to a scheme of Greek 
characters. The most consistent fissures form a series 
of mediolaterally oriented sulci designated a, (3 and 8, 
with other less prominent and less consistent sulci 
being present (Elliot Smith 1899, 1902; Hines 1929; 
Burkitt 1934; Lende 1964). The only departure from 
lissencephaly in the platypus cortex is the presence of 
a slight and shallow rhinal sulcus on the ventral surface 
of the cerebrum (Elliot Smith 1902). 

The divergence within monotreme evolution 
that has given rise to the lissencephalic and 
gyrencephalic forms of the cerebral cortex in the 
platypus and echidna is similar to that seen within both 
the placental and marsupial orders. Among the 
placental mammals, lissencephaly is apparent in the 
cortex of the rat and rabbit and even in some primates, 
such as the marmoset monkey (Callithrix jacchus; 
Brodmann 1909; Rowe et al.1996; Zhang et al. 1996, 
2001), whereas in the cat, the human being, and in a 



great many other placental mammals the cortex has 
acquired a prominent gyrencephalic form. However, 
the fact that the echidna and platypus display such 
striking differences in cortical folding serves as a clear 
reminder that the monotremes have had the same 
opportunity for evolutionary change and adaptation 
as have the species within the placental or marsupial 
orders. 

Quantitative indices of brain development in 
monotremes and other mammals 

A variety of quantitative measures of brain 
development also reinforce the conclusion that the 
brain of living monotremes is no more likely to provide 
a guide to the status of the ancestral mammalian brain 
than that of any contemporary placental mammal (for 
review, Rowe 1990). Furthermore, such measures fail 
to identify the living monotremes as being 
systematically less advanced in neurological terms than 
many of their marsupial and placental relatives. The 
quantitative measures invoked for these comparisons 
have been based usually upon indices related to the 
ratio of brain mass to total body mass and include, for 
example, the Encephalization Quotient {EQ) put 
forward by Jerison (1973) and defined for a given 
species as the ratio of actual brain mass (E { ) divided 
by the expected brain mass (E c ) for a mammalian 
species of a given body mass (P). The expected brain 
mass was obtained from the equation: 

E e = kPi 273 

that governs the overall relation between brain mass 
and body mass for the large range of living mammals 
for which Jerison gathered data. An EQ value of 1 
was assigned by Jerison for the average living placental 
mammal with values varying by a factor of 
approximately 25 times, from a low of -0.25 for basal 
insectivores, such as the hedgehog (Erinaceus 
europaeus) and the Madagascan tenrecs, through to 
values of ~6 - 7 for human beings and cetaceans such 
as the bottlenose dolphin (Tursiops truncatus). Values 
of -0.5 - 0.75 along the EQ scale were assigned to the 
echidnas by Jerison (1973) who argued that they '''are 
well in advance of didelphids like the opossum or 
insectivores like the hedgehog in relative brain size 
and differentation ". He argued that the monotremes 
should be considered, in terms of brain development, 
to be "at almost the same level as living progressive 
mammals" and speculated that they had reached this 
level by parallel evolution. 

It must be emphasized that comparisons of 
brain development based upon measures such as the 
EQ are somewhat arbitrary and that relativities arrived 



302 



Proc. Linn. Soc. N.S.W., 125, 2004 



M J. ROWE, D.A. MAHNS AND V. SAHAI 



at across species may change if other measures were 
to be used. However, a variety of alternative measures, 
including for example, the relative extent of the 
neocortex, once again fail to identify monotremes as 
being distinctly less developed neurologically than 
eutherian mammals (Pirlot and Nelson 1978; for 
review Rowe 1990). In addition, behavioural tests of 
discriminative learning in the echidna reveal an ability 
in particular, in spatial and positional discrimination 
tasks, that is not inferior to eutherian or metatherian 
species (Buchmann and Rhodes 1978). 



SENSORY AND PERCEPTUAL MECHANISMS 
IN MONOTREMES 

Behavioural observations from as early as the 
19 th century have pointed to the pre-eminence of the 
tactile sense in the perceptual life of both the platypus 
and the echidna (for review: Rowe 1990; Rowe et al. 
2003). In the case of the platypus it was apparent that, 
for both navigation and feeding, the animal relies, 
perhaps exclusively, upon sensory information from 
the bill as its eyes, nose and external auditory canals 
remain closed in the course of swimming and foraging 
(Bennett 1877; Burrell 1927; Griffiths 1978; Grant 
1984). While vision may be of more importance in the 
echidna than the platypus (see Gates 1978) it is 
probable that it assumes a lesser importance than the 
trigeminal tactile inputs from the snout and tongue 
which appear to play a similar prominent sensory role 
to that of the bill in the platypus. However, tactile 
information from the limb extremities, in particular, 
from the forelimb, also assumes great importance in 
the digging and burrowing activities of the echidna 
(Griffiths 1978; Augee and Gooden 1993). Because 
of this prominent sensory role for the distal forelimb 
as a tactile exploratory organ in the echidna we have 
recently undertaken an analysis of tactile neural 
mechanisms associated with the forepaw in 
Tachyglossus aculeatus (Manns et al. 2003; Rowe et 
al. 2003). As tactile neural mechanisms have been most 
intensively investigated for the distal glabrous skin of 
the forelimb in a great variety of species, this analysis 
permitted a comparison with placental mammals 
enabling us to establish the extent to which 
correspondence or divergence had arisen over the 
separate evolutionary paths taken within the different 
mammalian orders. 

Peripheral tactile neural mechanisms associated 
with the distal forelimb in the echidna: 
comparison with placental representatives 

Tactile sensory nerve fibres that arise from 



the distal glabrous skin of the limbs in the cat and in a 
variety of Old- and New-World monkeys fall into three 
major functional classes (Lindblom 1965; Lindblom 
and Lund 1966; Janig et al. 1968, Talbot et al. 1968, 
Iggo and Ogawa 1977, Ferrington and Rowe 1980; 
Ferrington et al. 1984; Coleman et al. 2001). These 
include one broad class that responds to static skin 
displacement with a so-called slowly adapting (SA) 
pattern of response that provides the basis for their 
designation as the SA class of fibres which, in both cat 
and primate species, appears to be associated with 
Merkel receptor endings (Janig et al. 1968, Talbot et 
al. 1968; Iggo and Muir 1969; Janig 1971; Iggo and 
Ogawa 1977; Ferrington and Rowe 1980; Coleman et 
al. 2001). 

The remaining tactile sensory nerve fibres 
supplying the primate hand or cat footpads are sensitive 
to only the dynamic components of tactile stimuli and 
can be divided into two distinct classes according to 
their sensitivity and responsiveness to cutaneous 
vibration (Janig et al. 1968; Talbot et al. 1968; Johnson 
1974; Iggo and Ogawa 1977; Ferrington and Rowe 
1980; Ferrington et al. 1984; Coleman et al. 2001). 
One class, designated the rapidly adapting (RA) or 
quickly adapting (QA) class is most sensitive to 
vibration at ~20-50Hz, and appears to be associated 
with intradermal, encapsulated receptors known as 
Krause corpuscles in the cat (Janig 1971; Iggo and 
Ogawa 1977) and as Meissner corpuscles in primates 
(Talbot et al. 1968; Coleman et al. 2001). The other 
purely dynamically-sensitive class (the PC class) is 
exquisitely sensitive to vibrotactile stimuli at 200- 
400Hz and is presumed to be associated with the 
Pacinian corpuscle (PC) class of receptor (Hunt 1960; 
Hunt and Mclntyre 1960; Sato 1961; Janig et al. 1968; 
Talbot et al. 1968; Lynn 1969; Ferrington and Rowe 
1980; Ferrington et al. 1984; Coleman et al. 2001). 

In human subjects where microneurography 
studies have permitted the recording and 
characterization of tactile sensory nerve fibres 
supplying the hand and finger tips, the same broad 
divisions apply, except that the SA group of fibres is 
reported to fall into two classes, designated the SAI 
fibres that appear to correspond with the single broad 
SA class in both the cat and non-human primates, and 
an SAII class that appears to be associated with Ruffini 
receptor endings, principally in the regions of skin 
around nail beds or skin creases near 
metacarpophalangeal joints (Knibestol and Vallbo 
1970; Johansson and Vallbo 1979). Although the SAII 
class is known to be present in the cat, it appears, in 
that case, to be confined to the hairy regions of skin, 
once again in association with Ruffini receptors 
(Chambers et al. 1972; Gynther et al. 1992). 



Proc. Linn. Soc. N.S.W., 125, 2004 



303 



MONOTREME TACTILE MECHANISMS 



100 pm 



200 pm 



400 pm 



600 pm 



1000 pm 



1200 pm 



1500 pm. 



4- 



* 



1 Second 



B 



■ n il i i ■! 1 1 i \\ 



800 pm III I II I I I I I I II . Illlll II 



60' 



40' 



20' 




T 



T 



L 



500 1000 

Step Amplitude (microns) 



1500 



Figure 1. Response traces and stimulus-response relations for representative low-thresholds SA afferent 
fibers supplying the glabrous skin of the echidna forepaw. (A): response traces to step indentations (lasting 
1.5s) at a range of amplitudes (100 to 1,500 um; represented in the two waveforms above and below the 
impulse traces). Stimuli were applied to the RF focus (indicated by the arrow in the inset in B) by means 
of a 2mm diameter stimulus probe. (B): stimulus-response relations for four SA fibers based on the mean 
response (impulses/sec ± S.D.) over the first one second from the onset of the 1.5s step indentation, plotted 
as a function of the indentation amplitude (modified from Mahns et al. 2003). 



For perhaps most mammalian species the 
densely innervated glabrous skin of the limb 
extremities represents the skin area of greatest tactile 
acuity (Darian-Smith 1984; Coleman et al. 2001; 
Johnson 2001, 2002) and may be regarded for most 
species as the tactile equivalent of the visual system's 
fovea. Some exceptions to this, where the tactile 
'foveal' role may be served by other structures, could 
include the bill in the platypus (Bohringer and Rowe 
1977; Rowe 1990; Rowe et al. 2003) and the nasal 
appendages of the star-nosed mole (Catania and Kaas 
1997). Nevertheless, it is probable that in the echidna, 
the distal forelimb glabrous skin assumes a similar 
crucial tactile sensory role as the footpads in the cat or 
the hand and fingers in primates. 

Electrophysiological analysis of tactile sensory 
nerve fibres supplying the echidna forepaw 

Microdissection of the ulnar or median nerves 
in the echidna forelimb permitted electrophysiological 
recording from almost 30 individual tactile sensory 
nerve fibres that supplied the glabrous regions of the 
echidna distal forelimb (Mahns et al. 2003). Once an 
individual sensory fibre was isolated, its cutaneous 
receptive field (RF) was mapped by means of gentle 
probing with von Frey hairs of known calibrated force. 
The functional characteristics of the fibre were then 



characterized by applying precise mechanical stimuli 
with a probe (usually 1-2 mm diameter) to the centre 
of the RF. The analysis, based upon the use of a 
feedback-controlled mechanical stimulator, revealed 
that the echidna tactile afferent fibres could be 
classified, like their placental counterparts, into two 
broad groups, one displaying slowly adapting (SA) 
response characteristics to static skin displacement, the 
other displaying a pure dynamic sensitivity. 

Functional characteristics of slowly adapting 
(SA) tactile afferent fibres supplying the echidna 
forepaw 

Tactile fibres in the SA class varied widely in 
their sensitivity to skin displacement. Those with 
lowest thresholds displayed small, circumscribed RFs 
and a sensitive grading of their impulse output as a 
function of skin displacement (Fig. 1A,B) and were 
therefore presumably well able to signal information 
about the location and intensity of skin perturbations 
encountered on the glabrous skin surface. A higher- 
threshold subclass of the SA fibres (Mahns et al. 2003) 
had larger RFs and less-well sustained responses 
(Fig.2A) and may contribute to the signalling of much 
more diffuse pressure encountered by the foot in 
locomotor or digging activity. The lesser sensitivity 
of this subgroup may be explained by the nature of 



304 



Proc. Linn. Soc. N.S.W., 125, 2004 



M.J. ROWE, D.A. MAHNS AND V. SAHAI 



B 



200 pm 
400 pm 
600 pm 
800 pm 
1000 pm 
1200 pm 
1500 pm 










° m m Mill I I I I I I I I I I II I I I L 


innn r mini in ii 1 1 1 1 1 1 1 1 1 1 1 1 1 mil 


1200um Hllll |j| || | I | I I | I I | | | | I I | I I |. 


1500um I IIIIIIIIIIIIIIIIIMI II I I I I I I I I I I Ii 




— *^ 1 second 1 — 



Figure 2. Response traces for a representative of the sensitive SA fibers associated with the echidna 
forepaw (A) and for claw-associated SA fibers. (B). The impulse traces in A and B show responses to the 
ramp indentation applied at a range of amplitudes (200-l,500um) to the glabrous skin (A) or to the base 
of the claw (B) (modified from Mahns et al. 2003). 



the echidna forepaw which is a very much thicker, 
cushioned structure than the footpad in the cat. This 
gross anatomical specialization appears to equip the 
echidna well for its robust digging and burrowing 
activity but may create, as a consequence of the 
viscoelastic properties of the footpad, a mechanical 
housing for many SA sensory endings that imposes 
limitations on the effective mechanical coupling 
between the skin surface and the receptor endings, and 
this, in turn, may account for the higher thresholds 
and more diffuse RF boundaries (Mahns et al. 2003). 

A further subset of SA fibres was associated 
with the base of the powerful claws and displayed a 
gradation in impulse output related to displacement at 
the base of the claw (Fig.2B), attributes that should 
equip them to subserve a kinaesthetic role in signalling 
movements of the powerful claws (Mahns et al. 2003). 

The low-threshold SA fibres appear to 
conform in properties to the SAI class of tactile afferent, 
already identified in the echidna snout (Iggo et al. 1996) 
and in the skin of placental species where it is generally 
thought to be associated with the Merkel receptor 
endings. Although the less-sensitive and claw- 
associated SA fibres in the echidna (Fig.2A,B) have 
some attributes resembling the placental SAII fibre 
class, in particular, a rather regular temporal pattern 
in the impulse activity of the claw-associated SA fibres 
(Mahns et al. 2003), the absence of spontaneous 
activity in these fibres is more consistent with an SAI 
identification. Thus, despite the rather disparate 
behaviour of the different subsets of SA fibres 
associated with the echidna forepaw it appears that, as 
a broad class, they probably conform more closely to 



the SAI class with its implied association with Merkel 
receptor endings. However, any conclusions about the 
receptor associations and precise identity if the echidna 
SA fibres must remain tentative until correlative 
histological data are available for the receptors. 

Functional characteristics of dynamically- 
sensitive tactile afferent fibres supplying the 
echidna forepaw 

Dynamically-sensitive tactile sensory fibres 
associated with the echidna forepaw could be divided 
into two classes principally according to their 
differential sensitivity to vibrotactile stimuli but, in 
addition, could often be distinguished on account of 
RF characteristics and sensitivity to manual stimuli 
(Mahns et al. 2003). One class resembled the rapidly 
adapting (RA) class of tactile afferent identified in 
association with the cat and primate glabrous skin in 
having small, circumscribed RFs and displaying 
maximum vibrotactile sensitivity at frequencies < 50 
Hz (Fig. 3). The second class had larger RFs and a 
bandwidth of vibrotactile sensitivity that extended up 
to 300-400 Hz, properties resembling those identified 
for the PC sensory fibres that are associated with 
Pacinian corpuscle (PC) receptors in the glabrous skin 
of placental species, including the cat, marmoset and 
macaque monkeys, and human subjects (reviewed 
above). 

Tactile coding capacities of dynamically-sensitive 
tactile afferent fibres in the echidna 

The RA class of afferents supplying the 
echidna footpad had absolute response thresholds of 



Proc. Linn. Soc. N.S.W., 125, 2004 



305 



MONOTREME TACTILE MECHANISMS 



B 



5 jam 



10 ^m 



20 urn 



40 urn 



100 um 




200 -i 



-o 150 


w 

"o. 

E 

^ 100 
en 

c 
o 

Q. 

(0 


* 50 



-I 




50 Hz 



T_ 



~ i i i 

50 100 150 200 

Vibration Amplitude (microns) 



Figure 3. Impulse records and stimulus-response relations for a representative RA afferent fiber supplying 
the echidna forepaw. A): Activation of the fiber as a function of amplitude increases in the 50Hz vibration 
train. The response achieves a 1:1 following pattern over the first ~15 cycles at 20 urn, and this response 
pattern is sustained throughout the 1 -second duration of the vibro tactile stimulus at 40 um. (B): Stimulus- 
response relations for this RA fiber show 1:1 plateau levels of response at <40 um at frequencies of 30 and 
50 Hz. At 100, 150, and 200 Hz, the 1:1 level was attained only at amplitudes of -70, 100, and 200 um, 
respectively. The RF for the fiber is indicated in the inset in B (modified from Mahns et al. 2003). 



<10 um at frequencies below -100 Hz. With increases 
in the intensity of vibrotactile stimuli, their response 
(in impulses/s) progressively increased until they 
attained a regular impulse discharge on successive 
cycles of the vibration stimulus. As the spike 
discharges in this so-called one-to-one pattern of 
response were tightly phaselocked to the vibration 
waveform the interspike intervals approximated the 
vibration cycle period. For example, at 50 Hz the 
interspike intervals approximated 20 ms and at 100 
Hz were ~10ms (Fig. 4). Because of the tight 
phaselocking, the pattern of discharge displayed a 
metronomic regularity that reflected very precisely in 
its temporal pattern the periodicity inherent in the 
vibrotactile stimuli. The impulse sequence thus 
provided a reliable signal, in an impulse pattern code, 
of the frequency, or pitch parameter of the vibrotactile 
stimuli. The tightness of phaselocking could be 
quantified by constructing cycle histograms (CHs; Fig. 
4B) which show the time of occurrence of impulses 
during each vibration cycle. The CHs use a pulse 



associated with the onset of each vibration cycle as a 
stimulus marker and have an analysis time that 
corresponds to the vibration cycle period. Responses 
that are tightly synchronized, or phaselocked, appear 
in the CH as a narrow peak as seen in Fig. 4B. When 
impulses occur independently of the phase of the 
applied vibration waveform the distribution in the 
histogram appears flat (Coleman et al. 2001; Mahns 
et al. 2003). The bandwidth of vibrotactile frequencies 
over which responses in the echidna RA fibres 
remained phaselocked extended from -5 Hz up to -200 
Hz, matching the behaviour of their placental 
counterparts in the cat (Ferrington and Rowe 1980; 
Ferrington et al. 1984; Rowe and Ferrington 1986) and 
in the macaque and marmoset monkeys (Talbot et al. 
1968; Coleman etal. 2001). 

Functional capacities of the presumed-PC class of 
tactile sensory fibre in the echidna 

Controlled vibrotactile stimuli permitted the 
distinction to be made between the RA afferent fibre 



306 



Proc. Linn. Soc. N.S.W., 125, 2004 



M.J. ROWE, D.A. MAHNS AND V. SAHAI 



50 HZ *" t ""^ ' " *'" w t ' ^M»r» " i» ^ ^^ 



U ww w t ' Un m*i mm W»»h Ww » im « Wnw#Jw 



B 




100 



Hz HHHHri444+f~ 



"2 ioo- 



o 
O 



100 Hz 
R = 0.94 




Figure 4. Precision of impulse patterning in the responses of echidna RA afferent fibers to vibrotactile 
stimuli. (A): Impulse traces show the tightly phase-locked pattern of response reflecting the periodicity of 
the 50 and 100 Hz vibrotactile stimuli. Quantitative measures of phase locking based on the vector strength 
or resultant, R (see Materials and Methods), were derived from the cycle histograms in (B), constructed 
to show the distribution of impulse occurrences throughout successive cycles of the vibration stimulus at 
the two frequencies. The analysis time in each CH corresponds to the cycle period of the vibration (modified 
from Mahns et al. 2003). 



class and a class of dynamically-sensitive tactile 
afferents with a distinctly broader bandwidth of 
sensitivity (Fig.5) reminiscent of placental PC fibres 
(Mahns et al. 2003). These putative PC fibres 
supplying the echidna forepaw had absolute response 
thresholds as low as ~5 [xm for the broad range of 
frequencies from -50 to 300 Hz (Fig.5C) which may 
equip the animal to detect small vibratory perturbations 
set up by termites in either the soil or in timber material 
encountered in the animal's use of the forepaw as an 
exploratory organ. Furthermore, these broad 
bandwidth vibrotactile sensors would be well-suited 
to serve as an early-warning system for the detection 
of ground-borne vibration signalling the movements 
of any predators or other animals in the vicinity 
(Mclntyre 1980; Mahns et al. 2003). As the PC fibre 
responses to vibrotactile stimuli remained tightly 
phaselocked at frequencies up to and beyond 400 Hz, 
the metronome-like patterning in their discharge (Fig. 
5A) would ensure that these fibres retained high acuity 
for signalling the temporal details of vibrotactile 
perturbations over a broad bandwidth of frequencies 
(Mahns et al. 2003). Although the bandwidth of 
vibrotactile responsiveness in the echidna PC fibres 
(Fig.5) did not extend to frequencies quite as high as 
those of placental PC fibres (Mahns et al. 2003) the 
explanation probably lies in the lower body 



temperature of the echidna (~28-32°C; Grigg et al. 
1992) rather than a fundamental difference in the 
receptors; in particular, as Sato (1961) has 
demonstrated that PC fibres in the cat display a 
displacement to lower frequencies in both bandwidth 
and peak sensitivity as temperature is lowered. 

The use of controlled sinusoidal vibration as 
a dynamic form of tactile stimulation in these studies 
permitted the precise quantification of both the 
frequency and intensive parameters of the stimuli. 
However, in addition, it provides a form of dynamic 
tactile stimulation that mimics in a controlled way the 
vibrational disturbances set up in the skin in association 
with relative movement between the skin and any 
textured surface encountered in the tactile exploratory 
movements of the forelimb. The properties revealed 
for echidna RA and PC fibres indicate that they are 
well suited to underpin the echidna's capacity to signal 
and code information about textural changes in the 
ground surface or in the coarseness or fineness of 
objects encountered, such as sand, gravel or soil, in 
the tasks of locomotion, digging and burrowing. 

In summary, it appears that for the distal 
forepaw of the echidna, the tasks of tactile exploration 
and perception are based upon a triad of major tactile 
sensory fibre classes, comprising a broad SA class and 
both RA and PC classes with functional capacities 



Proc. Linn. Soc. N.S.W., 125, 2004 



307 



MONOTREME TACTILE MECHANISMS 



A 

50 Hz 



C Q. 



444- 



1IHI I 



U 



I 



,MH '44444-M I It I I I H44444- 



C Q. 

31 

o 



ILL 



n 



It 



200 Hz 



-r- 



J2 „ 5 • 

C Q. 

2 E 



n 



W44rU44444^^ 



II I HI HI II 11 1 1 11 1 1 II 



400 Hz 



a„5 

C Q. 

<-> o 



1 1 ■ I J 1 1 ' 1 1 M I I 1 1 1 ! 



B 



400 Hz 



g> 300- 



200 



a) 100 





200Hz— 


300 Hz . 


. /-f/lOOHz, 


7 50 Hz 







10 Hz 



I ii I 



20 40 60 80 

Vibration Amplitude (microns) 



-. 50-, 

in 

c 

o 

I 40 
% 30 



20 



£ 10 



.......... 1:1 Tuning 

— «— Absolute 




100 



200 



300 



400 



500 



600 



Vibration Frequency (Hz) 



Figure 5. (A): Temporal patterning in the vibrotactile responses of PC -like fibers in the echidna. Impulse 
traces and peristimulus time histograms (PSTHs) show the metronome-like impulse pattern at the 1:1 
response level at vibrotactile frequencies of 50-400 Hz. Each PSTH was constructed from five consecutive 
responses to 20 cycles of vibration at each of the indicated frequencies. (B and C): Stimulus-response 
relations and vibrotactile frequency bandwidths for the putative PC class of echidna tactile afferent 
fiber. (B): Stimulus-response relations for a single PC -like fiber with RF on the lateral aspect of the 
forepaw glabrous skin, based on plots of the mean response (impulses/second) as a function of vibration 
amplitude at seven frequencies in the range 10-400 Hz. (C): Plots of absolute (solid lines) and 1:1 tuning 
thresholds (dashed lines) derived for five PC -like afferent fibers from stimulus-response data of the type 
shown in (B) (modified from Mahns et al. 2003). 



resembling those of the corresponding classes in 
placental mammals. The issue of whether the broad 
SA class might contain subsets will be resolved only 
with more detailed morpho-functional correlative 
analysis on both the receptor endings and the associated 
sensory nerve fibres. However, the breakdown of the 
echidna tactile sensory fibres into three broad classes 
resembling those in placental mammals suggests that 
peripheral mechanisms for tactile sensation in the distal 
glabrous skin are highly conserved across different 
mammalian orders (Mahns et al. 2003). 



CEREBRAL CORTICAL ORGANIZATION FOR 
TACTILE PROCESSING IN MONOTREMES 

The behavioural evidence for the pre- 
eminence of the tactile sense in the platypus, and 



probably also in the echidna, has been re-inforced by 
electrophysiological studies on the organization of the 
cerebral cortex in these two species. In the platypus in 
particular, the allocation of neocortical space to tactile 
processing is quite spectacular as was demonstrated 
with both evoked potential and single-neuron 
microelectrode recording studies first undertaken in 
our laboratory in the 1970s (Bohringer and Rowe 1977; 
Rowe 1990), and more recently by Krubitzer et al. 
(1995a). With evoked potential mapping, a brief 
electrical stimulus delivered at a point on the skin 
surface activates sensory fibres that generate a 
synchronous input to the areas of cerebral cortex 
involved in processing information from that source 
of sensory input. From the cortical surface overlying 
these areas it is possible to record an evoked potential 
that is usually biphasic, consisting of an initial positive- 
going deflection followed by a larger negative-going 



308 



Proc. Linn. Soc. N.S.W., 125, 2004 



M.J. ROWE, D.A. MAHNS AND V. SAHAI 




200mV 



Figure 6. Evoked potentials (positively downwards) recorded from the cortical surface of the platypus 
following bipolar stimulation of the anterolateral margin of the contralateral bill (IV, 100-microseconds 
pulse). Each recording was made from the position indicated by the dot at the left of the trace. Dotted 
lines indicate positions of large blood vessels in frontal region of hemisphere; view of hemisphere and 
whole brain (inset) from dorsolateral aspect. Stippled areas in inset represent focal projection sites and 
include sites at which positive-going responses exceed lOOuY for bill (B), 30uV for fore limb (FL) and 
lOuV for hind limb (HL). Zones between stippling and continuous lines include sites from which smaller 
responses could be recorded (from Bohringer and Rowe 1977). 



component (Fig.6, and Bohringer and Rowe 1977). 
The initial positive-going component is thought to arise 
from the direct excitatory action of thalamo-cortical 
afferent input on cortical neurons (Mountcastle and 
Poggio 1968), and therefore the cortical region from 
which this component can be recorded is thought to 
represent the projection focus for that source of input. 
The evoked potentials illustrated in Fig.6 were 
recorded in response to stimulation on the anterolateral 
margin of the bill and reveal that a vast area of the 
dorsal surface of the contralateral cerebral hemisphere 
is taken up with the processing of bill inputs. At each 
recording point indicated by the dots on the main 
figure, responses were also recorded to forelimb (FL) 
and hindlimb (HL) stimulation in the platypus allowing 
the isopotential contour maps for these and the bill 



(B) inputs to be constructed on the inset figure, with 
the stippled areas indicating the focal projection sites 
for the three sources of input, and the zone between 
the stippling and continuous line a region from which 
smaller evoked potentials could be recorded. 
Single neuron mapping of the somatosensory 
cortex in monotremes 

More detailed single-neuron microelectrode 
recordings from as many as 250 individually- 
discriminated cortical neurons in up to 67 electrode 
penetrations in a given experiment on the platypus 
somatosensory cortex confirmed the highly ordered 
and complete cortical representation of tactile inputs 
from the contralateral body, extending from the mid- 
sagittal region of the hemisphere out to the region of 
the rhinal sulcus (Figs.7 and 8) on the ventrolateral 



Proc. Linn. Soc. N.S.W., 125, 2004 



309 



MONOTREME TACTILE MECHANISMS 



surface, the one sulcus present as an exception to the 
lissencephalic state of the platypus cerebral cortex. As 
the microelectrode mappings were based on inputs 
generated by light tactile stimulation of the skin surface 
they confirm the remarkable size of the cortical space 
devoted to tactile processing, in particular, from the 
bill. This is important as the electrical stimuli delivered 
to the skin in evoked potential mappings could have 
activated a combination of tactile and the putative 
electroreceptive afferent fibres postulated to be present 
in the bill of the platypus (see below). The continuity 
of tactile representation across the somatosensory 
cortex of the platypus was confirmed in several detailed 
mappings (Bohringer and Rowe 1977) and is apparent 
in Fig. 8 which shows the tactile receptive fields 
outlined on the figurines for 60 individual neurons 
isolated in nine electrode penetrations made in a single 
anteroposterior plane in the posterior region of the 
hemisphere. While tactile receptive fields for 
individual neurons are up to -15 cm 2 on regions such 
as the tail and trunk, those on the distal glabrous skin 
of the limbs were much smaller, while those on the 
bill, in particular, its anterior and lateral margins, were 
no more than 1mm in diameter. These represent the 



smallest tactile receptive fields ever recorded in the 
cortex and are therefore capable of conferring great 
precision and fidelity upon the tasks of tactile 
localization and discrimination involving the bill. 

Where the electrode penetration was made 
normal to the cortical surface, the neurons encountered 
had very similar receptive field locations (Fig. 8) 
indicative of the columnar organization, well described 
for the cerebral cortex of placental mammals (e.g. 
Mountcastle 1957), in which neurons of similar 
functional properties are grouped in columns oriented 
normal to the surface. In penetrations, such as number 
eight in Fig.8, that passed obliquely through successive 
cortical columns in the region of bill representation, 
there was a remarkably orderly and progressive shift 
in the representation of the bill surface that is apparent 
in the enlarged and expanded view of this electrode 
track in Fig. 9, emphasizing once again the striking, 
fine-grain spatial resolution available within the area 
of bill representation in the somatosensory cortex of 
the platypus. 

The somatosensory cortex of the echidna, 
mapped by Lende (1964), also has a striking allocation 
of space to the tongue and snout representations but, 



Figure 7. Inset shows entry points (filled circles) for 54 microelectrode penetrations into the platypus 
cortex. The black areas on figurines show the combined receptive field areas for all neurons (up to 15) 
sampled in each of the penetrations. Dotted lines represent positions of large blood vessels in frontal 
region of hemisphere; view of hemisphere from dorsolateral aspect (from Bohringer and Rowe 1977). 




"llrW t r' y <?' y 

i>i»t 

^^0000 
\ \q ixsOOOOQ 

boo odd ' 
ooooo 
oooo 




310 



Proc. Linn. Soc. N.S.W., 125, 2004 



M.J. ROWE, D.A. MAHNS AND V. SAHAI 




Figure 8. (A): Photograph of cortical surface of the platypus from dorsolateral aspect indicating a plane 
in which 9 penetrations were made. (B): Coronal section through hemisphere at the plane indicated in A, 
showing the course of the 9 penetrations. (C): Reconstruction of penetrations 1-9 indicating location of 
each neuron studied (filled circles) and its peripheral receptive field. No fields could be found for the first 
two neurons in penetration 4. Receptive fields for all neurons in each of the penetrations 7-9 were confined 
to the shaded areas on each of the associated figurines (from Bohringer and Rowe 1977). 



in addition, a prominent representation of the distal 
forearm, presumably reflecting its importance in 
digging and burrowing. 

Multiple representation of the body within the 
monotreme cerebral cortex 

For both the platypus and the echidna, the 
early cortical mapping studies in our laboratory 
(Bohringer and Rowe 1977; Rowe 1990) and that of 
Lende (1964), led to the conclusion that there was a 
single body representation in the contralateral cerebral 
hemisphere, conforming to the so-called primary 
somatosensory cortex (SI) of other mammals. No 
evidence was found for a second representation that 



might correspond to either the SII area that is well 
recognized in, for example, the cat and primate species 
(for review, Rowe 1990; Johnson 1990; Rowe et al. 
1996; Zhang et al. 1996, 2001) or indeed, to any other 
areas of somatosensory representation that have been 
reported in some placental species (Kaas 1982, 1987; 
Krubitzer and Kaas 1990; Krubitzer et al. 1995b), and 
now more recently, for both the echidna and platypus 
(Krubitzer et al. 1995a). Krubitzer et al. (1995a) have 
proposed that there are four somatosensory 
representations within the contralateral cerebral cortex 
of both the platypus and the echidna which they 
designate the primary somatosensory cortex (57), the 
Rostral deep field (R), the Manipulation field (M) and 



Proc. Linn. Soc. N.S.W., 125, 2004 



311 



MONOTREME TACTILE MECHANISMS 




Figure 9. Coronal section of platypus cerebral cortex showing the 
position of a microelectrode penetration (track 8 in Fig.8) made 
obliquely to the cortical surface. The filled circles indicate the locations 
of the 14 neurons studied. The receptive field for each neuron is 
indicated on the right which is an enlargement of the area on the 
ventral surface of the bill (modified from Bohringer and Rowe 1977). 



the Parietal Ventral field (PV). Furthermore, each is 
said to contain "a complete representation of the body 
surface", a surprising claim to us, considering that the 
four areas are not all defined as being concerned with 
the processing of mechanosensory data from the body 
surface. The area given the SI designation is concerned 
principally with cutaneous inputs, whereas area R 
immediately rostral to SI is said to contain neurons 
that respond "most often to stimulation of deep 
receptors" and which required light taps or light 
pressure to the body surface to elicit a response. 
However, as the tactile stimuli employed in their study, 
whether brushing, tapping, or pressure forms of 
cutaneous stimulation, were neither quantified nor 
reproducible, it is difficult to see how reliable 
distinctions were made in terms of neural response 
thresholds between neurons in the different cortical 
representation areas. 

Topographic organization of the putative 
multiple areas of somatosensory representation in 
the monotreme cerebral cortex 

As responsiveness criteria may be insufficient 
to permit an unequivocal division of the monotreme 
somatosensory cortex into four distinct areas 
(Krubitzer et al. 1995a), such a differentation must 
therefore depend upon topographic considerations, in 
particular upon the extent to which the four putative 
areas constitute complete and discrete representational 
maps of the body. In the case of adjacent areas, 



whether, for example, SI and 
R, or SI and PV, Krubitzer et 
al. (1995a) state that the 
boundaries coincide with a 
reversal in the representation 
of peripheral receptive fields as 
one progresses across a 
sequence of cortical recording 
sites. For example, in both 
platypus and echidna cortex, 
the receptive fields on the 
upper body may shift from 
proximal body locations, 
including the face, shoulder 
and upper limb, to the distal 
forelimb, and then move back 
to more proximal parts of the 
limb and shoulder. Similar 
reversals were seen across 
sequences of recording sites 
involving the hindlimb 
representations, leading to the 
interpretation by Krubitzer et 
al. (1995a) that the separate 
representations of proximal limb and associated trunk 
regions constitute parts of two distinct body 
representations. However, an alternative interpretation 
must be considered, which emerges from the detailed 
studies carried out in the macaque monkey by Werner 
and Whitsel (1968, 1973) and Whitsel et al. (1969, 
1971) for SI in the postcentral gyrus, and for the more 
laterally-placed second somatosensory area, SII. 

The representation of the body within the 
cerebral cortex: a reflection of the dermatomal 
trajectory 

The crucial observation emphasized by 
Werner and Whitsel was that the body maps in both SI 
and SII of the cerebral cortex owe their essential 
topographic properties to the serial and overlapping 
projection of the dorsal roots. In the case of the 
macaque postcentral gyrus, the representation of spinal 
roots, from sacral through to cervical, forms a 
succession of antero-posteriorly oriented bands 
progressing from medial to lateral across the cortex 
(e.g., Figs. 6 and 8 in Werner and Whitsel 1973). 

Within the SI area of the macaque monkey, 
and indeed within the macaque SII as well, the 
representations of the postaxial and preaxial arm and 
leg areas are separated by the representation of the 
more distal parts of the limbs, in particular, the digits 
and toes respectively. This effectively gives rise to a 
split representation of the upper parts of the limbs. In 
the case of the postcentral gyrus, one component lies 



312 



Proc. Linn. Soc. N.S.W., 125, 2004 



M.J. ROWE, D.A. MAHNS AND V. SAHAI 



medial to the distal limb representation, the other lateral 
to it, an arrangement represented schematically to 
illustrate this dermatomal trajectory in Fig. 8 of Werner 
and Whitsel (1973). One may observe how the 
dermatomal trajectory generates the split representation 
of the upper regions of either forelimb or hindlimb 
within the cerebral cortex by examining, in a human 
anatomy text (e.g., Williams and Warwick 1980) the 
dermatomal boundaries of successive spinal roots 
associated with either the lower or upper regions of 
the body. In the case of the upper body, the ventral 
and dorsal axial lines of the upper limb mark a border 
between the innervation fields of C5 and T 1 . Therefore, 
in any representation of the body surface within the 
cerebral cortex, one might expect, with the central map 
being laid down according to the dermatomal trajectory 
(Werner and Whitsel 1968, 1973; Whitsel et al. 1969, 
1971), that the central representation of the upper arm 
will be split along the axial lines with the input from 
the distal limb, carried over the C6, 7 and 8 roots, 
creating a clear separation in the cortical representation 
of lateral and medial surfaces of the upper arm. 

As the same fundamental plan and sequence 
for tactile dermatomes is also found in both the cat 
and monkey (Sherrington 1898; Kuhn 1953; 
Hekmatpanah 1961), one may assume that this 
organizational plan for spinal segmental innervation 
is a general one that would operate across mammalian 
orders, including the monotreme representatives. 

Is there multiple representation of the body 
within the platypus and echidna cerebral cortex? 

If one examines the receptive fields plotted 
for the platypus and echidna somatosensory cortex by 
Krubitzer et al. (1995a) in their Figs. 6 and 16, it might 
be argued that those fields on the proximal parts of the 
limb, on either side of the distal limb representation, 
are not clearly and systematically separated into 
representations of the medial and the lateral surfaces 
of the limb as might be expected in a perfect reflection 
of the dermatomal trajectory. However, this is hardly 
surprising on several grounds that are outlined in a 
recent review (Rowe, in press). 

The fundamental point to be emphasized is 
that the reversals in receptive field representation 
described by Krubitzer et al. are consistent with the 
sequence of representation that might be expected 
within a single body map whose plan is determined 
by the dermatomal trajectory. In view of these 
considerations we would re-emphasize our 1977 
finding (Bohringer and Rowe 1977; Rowe 1990) of a 
single large representation of the contralateral body 
within the platypus cerebral cortex and the similar 
conclusion reached even earlier by Lende (1964) for 



the echidna. Furthermore, we wish to emphasize the 
fundamental importance and significance of Werner 
and Whitsel's studies (1968, 1973; and Whitsel et al. 
1969, 1971) on body representation within the cerebral 
cortex, and the need that arises from their studies, to 
take account of the dermatomal trajectory as the crucial 
determinant of representational topography within 
central neural systems. 

Sensory and perceptual specialization in 
monotremes: trigeminal electroreceptive 
mechanisms 

As the monotremes emerged in mammalian 
evolution on a separate line from therian mammals in 
the early Mesozoic (Dawson 1983; Rowe 1990; Augee 
and Gooden 1993) the possibility that some 
qualitatively different apparatus for neural sensing 
might have emerged was given some credence in the 
1980s with both behavioural and electrophysiological 
studies reporting the presence of electroreception in 
monotremes (Scheich et al. 1986; Gregory et al. 1987, 
1988, 1989a, b). These and subsequent reports 
suggested that electroreception might be associated 
with the bill of the platypus and the snout of the 
echidna, in each case in association with the trigeminal 
nerve rather than the lateral line system that is the 
principal basis of electroreception in certain fish (Cahn 
1967; Bullock 1999). However, neither behavioural 
nor electrophysiological data have provided any 
suggestion of electroreception in association with other 
skin regions or somatosensory nerves of the 
monotremes, such as the median or ulnar nerves of 
the forelimb. 

The first behavioural evidence for 
electroreception in the platypus bill came in a short 
report from Scheich et al. ( 1 986) that the platypus could 
detect weak electric fields with threshold strengths as 
low as 50-200 \iV cm 1 . Furthermore, they reported 
that an electroreceptive processing zone was present 
in the posterolateral region of the contralateral cerebral 
cortex on the caudal side, but next to, the map of bill 
mechanoreceptive input identified by Bohringer and 
Rowe (1977). In our view this is a puzzling finding on 
several grounds (for review, see Rowe 1990). First, in 
an earlier detailed electrophysiological mapping of the 
cerebral cortex we found that the cortical region 
concerned with tactile representation of the bill 
occupied almost the whole of the posterolateral region 
of cortex (Bohringer and Rowe 1977; Rowe 1990). 
Furthermore, we constructed separate cortical maps 
of bill representation based upon mechanical 
stimulation which was specific for tactile inputs, and 
electrical stimulation which would have activated both 
tactile and the putative electroreceptive afferents. 



Proc. Linn. Soc. N.S.W., 125, 2004 



313 



MONOTREME TACTILE MECHANISMS 



However, comparison of the two maps (Figs.6 and 7; 
and Bohringer and Rowe 1977) reveals no evidence 
of a separate area of bill representation in the map based 
upon the electrical stimulation, from that obtained with 
pure tactile stimulation. In contrast to the Scheich et 
al. (1986) report that electroreceptive-induced cortical 
evoked potentials were found next to the map of bill 
mechanoreceptor input, those by Iggo et al. (1992) and 
Krubitzer et al. (1995a) indicated that the 
electrosensory area of cortical representation lay 
entirely within the border of the tactile representation 
area defined for the bill in our earlier study (Bohringer 
and Rowe 1977; Rowe 1990). However, Krubitzer et 
al. (1995a) have described a "clear functional 
parcellation" within this SI region whereby regions 
responsive to just tactile input were interdigitated with 
regions responsive to both tactile and electroreceptive 
inputs (e.g. Fig. 10 in Krubitzer et al. 1995a). 
Furthermore, the regions of pure tactile sensitivity 
coincided with myelin and cytochrome oxidase-dense 
regions of SI while the regions of putative bimodal 
sensitivity coincided with myelin-light regions. What 
the significance of such differences in myelin density 
might be in this circumstance remains unclear. 
However, there may not be universal agreement with 
this assertion anyway, judging by the allocation of light 
and dark areas in their Fig.lOB; for example, the 
myelin-dark region drawn to contain four recording 
sites of pure mechanosensitivity in part B of their figure 
appears rather paler in the adjacent tangentially- 
oriented photomicrograph than some areas represented 
as myelin-light in Fig.lOB. It is likely to be difficult to 
make these distinctions reliably, in the tangentially- 
cut cortical sections, firstly without objective analysis 
of image density, and secondly, in the absence of 
systematic control for laminar depth across the extent 
of the cortical section under study. 

Electrosensory field-strength thresholds 
illustrated for cortical neurons in Fig. 10 by Krubitzer 
et al. (1995a) ranged up to 900 uV cm -1 ; however, it 
was not clear what field strength might have activated 
neurons in the purely tactile-sensitive regions. 

Behavioural and afferent fibre thresholds for 
electroreception 

A further concern in relation to claims for 
electroreception in monotremes arises over the 
thresholds reported for the phenomenon. At the 
behavioural level, Scheich et al. (1986) reported field 
strength values as low -50-200 u.V cm' 1 for the 
platypus, and, more recently, values of 20uVcm~' have 
been reported (Manger and Pettigrew 1996; Pettigrew 
1999). However, individual trigeminal afferents, 
believed to be of the electrosensitive class, had 



threshold field strengths of ~4mV cm 1 (Gregory et al. 
1988, 1989b). As these values were based on a 
substantial fibre sample, and as these values are vastly 
higher than reported behavioural thresholds for the 
platypus, one might infer that any putative 
electroreceptive sense must depend upon some other 
source of afferent input. Proposals that spatial 
summation, based upon convergence of a number of 
trigeminal electroreceptive afferents onto central 
neurons, may confer the observed behavioural 
thresholds upon the animal are difficult to accept when 
none of the sampled afferent fibres has thresholds low 
enough to account for this. 

To the extent that there are contentious issues 
associated with the claims for electroception within 
the monotreme order of mammals, it is important that 
further rigorously controlled investigations be pursued 
to resolve such issues, in particular, more detailed 
behavioural studies based upon objective analysis 
rather than anecdotal accounts of the movement 
patterns of the platypus and echidna in relation to 
presumed electrosensory stimuli. 



REFERENCES 

Augee, M. and Gooden, B. (1993). 'Echidnas of Australia 

and New Guinea'. Sydney, University of New 

South Wales Press. 
Belov, K. and Hellman, L. (2003). Platypus 

immunoglobulin M and the divergence of the 

two extant monotreme lineages. Australian 

Mammalogy 25, 87-94. 
Bennett, G.F. (1877). Notes on Ornithorhynchus 

paradoxus. Proceedings of the Zoological 

Society (Lond.) 

161-166. 
Bohringer, R.C. and Rowe, M.J. (1977). The organization 

of the sensory and motor areas of cerebral cortex 

in the Platypus (Ornithorhynchus anatinus). 

Journal of Comparative Neurology 174, 1-14. 
Brodmann, K. (1909). l Vergleichende Lokalissationslehre 

der Grosshirnrinde '. (Barth: Leipzig). 
Buchmann, O.L.K. and Rhodes, J. (1978). Instrumental 

learning in the echidna Tachyglossus aculeatus 

setosus. The Australian Zoologist 20, 131-145. 
Bullock T.H. (1999). The future of research on 

electroreception and electrocommunication. 

Journal of Experimental Biology 202, 1455- 

1458. 
Burkitt, A.N.S. (1934). The variability of the gyri and 

sulci in the cerebral hemispheres of 

Tachyglossus (echidna) aculeata, Psychiatria et 

neurologia 38, 368-378. 
Burrell, H. (1927). 'The Platypus: Its Discovery, 

Zoological Position, Form and Characteristics, 

Habits, Life History Etc'. (Angus and 

Robertson: Sydney). 



314 



Proc. Linn. Soc. N.S.W., 125, 2004 



M.J. ROWE, D.A. MAHNS AND V. SAHAI 



Cahn, P.H. (1967). 'Lateral line detectors'. (Indiana 

University Press: Bloomington). 
Catania, K.C. and Kaas, J.H. (1997). Somatosensory fovea 

in the star-nosed mole: behavioral use of the star 

in relation to innervation patterns and cortical 

representation. Journal of Comparative 

Neurology 387, 215-283. 
Chambers, M.R., Andres, K.H., von During, M. and Iggo, 

A. (1972). The structure and function of the 

slowly adapting type II mechanoreceptor in 

hairy skin. Quarterly Journal of Experimental 

Physiology 57, 417-445. 
Coleman, G.T., Bahramali, H., Zhang, H.Q. and Rowe, 

M.J. (2001). Characterization of tactile afferent 

fibers in the hand of the marmoset monkey. 

Journal of Neurophysiology 85, 1793-1804. 
Dawson, T.J. (1983). 'Monotremes and Marsupials: The 

Other Mammals'. (Arnold: London). 
Darian-Smith, I. (1984). The sense of touch: performance 

and peripheral neural processes. In 'Handbook 

of physiology the nervous system III' (Eds J.M. 

Brookhart, V.B. Mountcastle, I. Darian-Smith 

and S.R. Geiger) pp.739-788. (American 

Physiological Society: Bethesda, MD). 
Elliot Smith, G. (1899). Further observations on the 

anatomy of the brain in the Monotremata. 

Journal of Anatomy and Physiology (Lond.) 33, 

309-344. 
Elliot Smith, G. (1902). Descriptive and illustrated 

catalogue of the physiological series of 

comparative anatomy, In 'Royal College of 

Surgeons Museum Catalogue of Physiology'. 

Volume 2, 2 nd (Ed Taylor and Francis) London. 
Ferrington, D.G. and Rowe, M.J. (1980). Functional 

capacities of tactile afferent fibres in neonatal 

kittens. Journal of Physiology (Lond.) 307, 335- 

353. 
Ferrington, D.G., Hora, M.O.H. and Rowe, M.J. (1984). 

Functional maturation of tactile sensory fibers in 

the kitten. Journal of Neurophysiology 52, 74- 

85. 
Gates, G.R. (1978). Vision in the monotreme echidna 

(Tachyglossus aculeatus). The Australian 

Zoologist 20, 147-169. 
Grant, T. (1984). 'The Platypus' (New South Wales 

University Press: Sydney). 
Griffiths, M. (1978). 'The Biology of the Monotremes' 

(Academic Press: New York). 
Gregory, J.E., Iggo, A., Mclntyre, A.K. and Proske, U. 

(1987). Electroreceptors in the platypus. Nature 

326, 386-388. 
Gregory, J.E., Iggo, A., Mclntyre, A.K. and Proske, U. 

(1988). Receptors in the bill of the platypus. 

Journal of Physiology (Lond.) 400, 349-366. 
Gregory, J.E., Iggo, A., Mclntyre, A.K. and Proske, U. 

(1989a). Responses of electroreceptors in the 

snout of the echidna. Journal of Physiology 

(Lond.) 414, 521-538. 
Gregory, J.E., Iggo, A., Mclntyre, A.K. and Proske, U. 

(1989b). Responses of electroreceptors in the 



platypus bill to steady and alternating potentials. 

Journal of Physiology (Lond.) 408, 391-404. 
Grigg, G, Augee, M.L. and Beard, L. (1992). Thermal 

relations of free-living echidnas during activity 

and in hibernation in a cold climate. In 'Platypus 

and echidnas' (Ed M.L. Augee) pp. 160-173. 

(Sydney: Royal Zoological Society: New South 

Wales). 
Gynther, B.D., Vickery, R.M. and Rowe, M.J. (1992). 

Responses of slowly adapting type II afferent 

fibres in cat hairy skin to vibrotactile stimuli. 

Journal of Physiology (Lond.) 458, 151-169. 
Hekmatpanah, J. (1961). Organization of tactile 

dermatomes, C ( through L 4 , in the cat. Journal 

of Neurophysiology 24, 124-140. 
Hines, M. (1929). The brain of Ornithorhynchus anatinus, 

Philosophical Transactions of the Royal Society 

of London Series B 217, 155-259. 
Hunt, C.C. (1960). On the nature of vibration receptors in 

the hindlimb of the cat. Journal of Physiology 

(Lond.) 155, 175-186. 
Hunt, C.C. and Mclntyre, A.K. (1960). Characteristics of 

responses from receptors from the flexor longus 

digitorum muscle and the adjoining interosseous 

region of the cat. Journal of Physiology (Lond.) 

153, 74-87. 
Iggo, A. and Muir, A.R. (1969). The structure and 

function of a slowly adapting touch corpuscle in 

hairy skin. Journal of Physiology (Lond.) 200, 

763-796. 
Iggo, A. and Ogawa, H. (1977). Correlative physiological 

and morphological studies of rapidly adapting 

mechanoreceptors in cat's glabrous skin. 

Journal of Physiology (Lond.) 266, 275-296. 
Iggo, A., Gregory, J.E. and Proske, U. (1996). Studies of 

mechanoreceptors in skin of the snout of the 

echidna Tachyglossus aculeatus. Somatosensory 

and Motor Research 13, 129-138. 
Iggo, A., Gregory, J.E. and Proske, U. (1992). The central 

projection of electrosensory information in the 

platypus. Journal of Physiology 447, 449-465. 
Janig, W., Schmidt, R.F. and Zimmermann, M. (1968). 

Single unit responses and the total afferent 

outflow from the cat's foot pad upon mechanical 

stimulation. Experimental Brain Research 6, 

100-115. 
Janig, W. (1971). Morphology of rapidly and slowly 

adapting mechanoreceptors in the hairless skin 

of the cat's hind foot. Brain Research 28, 217- 

231. 
Jerison, H.J. (1973). 'Evolution of the Brain and 

Intelligence' (Academic Press: New York). 
Johansson, R.S. and Vallbo, A.B. (1979). Tactile 

sensibility in the human hand: relative and 

absolute density of four types of 

mechanoreceptive units in glabrous skin. 

Journal of Physiology (Lond.) 286, 283-300. 
Johnson, J.I. (19